Neuropsychology © 2014 American Psychological Association

2015, Vol. 29, No. 3, 463– 472 0894-4105/15/$12.00 http://dx.doi.org/10.1037/neu0000155

Functional and Structural Indices of Empathy: Evidence for

Self-Orientation as a Neuropsychological Foundation of Empathy
Brick Johnstone, Dan Cohen, Kirk R. Bryant, Bret Glass, and Shawn E. Christ
University of Missouri

Objective: To evaluate a model that hypothesizes that empathy is associated with decreased right parietal
lobe (RPL)-related self-orientation (i.e., increased selflessness), which allows individuals to more easily
empathize with others. Methods: Participants: Thirty one individuals with documented neuroradiological
abnormalities due to traumatic brain injury (TBI) referred for clinical evaluations. Measures: Cerebral
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.

integrity was measured with both functional (i.e., neuropsychological tests) and structural indices (i.e.,
This document is copyrighted by the American Psychological Association or one of its allied publishers.

MRI). Participants were administered comprehensive neuropsychological tests associated with general
bilateral frontal, temporal, and parietal lobe functioning, a self-report measure of empathy (i.e., Penner’s
Prosocial Personality Battery), and an objective measure of empathy (i.e., Prisoner’s Dilemma). Twenty
participants also completed structural MRI analysis of the bilateral frontal, temporal, parietal, and insular
cortices measured in terms of volume. Results: Pearson correlations indicated that empathy was related
to increased neuropsychological indices of RPL and frontal lobe (primarily left frontal) functioning. The
only MRI indices associated with empathy were the bilateral insula. Neither functional nor structural
cerebral indices were significantly related to objective measures of empathy. Conclusions: Contrary to
hypotheses, empathy appears to be associated with increased RPL functioning. It is suggested that to
incorporate the experiences of others into the experience of the self (i.e., to be empathetic), one must have
an intact sense of the self.

Keywords: affective empathy, cognitive empathy, self-orientation, right parietal lobe, neuropsychology

Neuroanatomy of Empathy
There has been growing interest in determining the neuroanat-
omical and neuropsychological foundations of empathy (hereby
defined as “the action of understanding, being aware of, being
sensitive to, and vicariously experiencing the feelings, thoughts,
and experience of another”). A meta-analysis of empathy research
concluded that empathy is a complex construct that is related to
multiple neurologic networks and cognitive and emotional abilities
(Zaki & Ochsner, 2012). Specifically, empathy has been shown to
be related to the inferior parietal lobe, temporoparietal junction,
anterior insula, posterior superior temporal sulcus, temporal pole,
premotor cortex, posterior cingulate cortex, anterior cingulate cor-
tex, medial prefrontal cortex, insula, right temporoparietal region,
and various regions of the frontal lobes (Banissy, Kanai, Walsh, &
Rees, 2012; Decety & Jackson, 2004; Decety & Jackson, 2006;
Eslinger, 1998; Grattan, Bloomer, Archambault, & Eslinger, 1994;
Grattan & Eslinger, 1989; Grattan & Eslinger, 1992; Mutschler,
Reinbold, Wankerl, Seifritz, & Ball, 2013; Preston & de Waal,
2002; Shamay-Tsoory, Tomer, Goldsher, Berger, & Aharon-
This article was published Online First November 17, 2014.
Brick Johnstone, Department of Health Psychology, University of Missouri;
Dan Cohen, Department of Religious Studies, University of Missouri; Kirk R.
Bryant, Department of Health Psychology, University of Missouri; Bret Glass,
Department of Psychological Sciences, University of Missouri; Shawn E.
Christ, Department of Psychological Sciences, University of Missouri.
Correspondence concerning this article should be addressed to Brick John-
stone, Department of Health Psychology, University of Missouri, DC116.88,
Columbia, MO 65212. E-mail: johnstoneg@health.missouri.edu
463
Peretz, 2004; Zaki & Ochsner, 2012). Considered together, these
studies implicate most of the brain in empathy, which is not
surprising given its complexity. This variability in findings is
likely related to the lack of clear definitions of empathy, method-
ological weaknesses inherent in neuroradiological scanning (Logo-
thetis, 2008), and other methodological issues (e.g., use of self-
report vs. objective measures of empathy; “artificiality” of
objective empathy measures; structural vs. functional measures of
cerebral integrity; use of clinical and nonclinical samples; Zaki &
Ochsner, 2012).
The majority of research on the neurology of empathy has
involved neuroradiological evaluation of nonclinical populations
engaged in objective computer-based tasks. However, determining
associations between empathy and cerebral regions is complicated
given the numerous other cognitive processes that are engaged
during objective tasks of empathy (e.g., attending to stimuli, lis-
tening to instructions, perceiving stimuli, cognitively processing
economic games commonly used in empathy research, etc.; Logo-
thetis, 2008). As a result, the neuroanatomical basis of empathy
has been difficult to determine. An alternative research method has
involved the investigation of populations with brain dysfunction in
which persons with various lesions/disorders are compared with
one another (e.g., persons with left vs. right frontal lobe lesions),
or to healthy control groups, in terms of empathy (Beadle &
Tranel, 2013; de Sousa et al., 2010; Eslinger, 1998; Grattan et al.,
1994; Grattan & Eslinger, 1989; Grattan & Eslinger, 1992; Rankin,
Kramer, & Miller, 2005; Rankin et al., 2006; Shamay-Tsoory et
al., 2004; Shamay-Tsoory, Aharon-Peretz, & Perry, 2009). Using
this method, empathy can be associated with different cerebral
areas (i.e., damage to areas associated with empathy will be
 464 JOHNSTONE, COHEN, BRYANT, GLASS, AND CHRIST
indicative of decreased empathy). However, a weakness in such
research is that only general relationships can be inferred given the
diffuse nature of most brain disorders. Whereas the first method
may lead to overidentification of the cerebral areas/functions spe-
cifically associated with empathy, the latter one can only be used
to make general inferences regarding brain-empathy relationships.
In fact, it has been suggested that the two methodologies are
appropriate complements to one another (Rorden & Karnath,
2004).
Neuropsychology of Empathy
In addition to identifying the cerebral areas associated with
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.
empathy, research has also focused on determining the specific
This document is copyrighted by the American Psychological Association or one of its allied publishers.
affective and cognitive components of empathy. Empathy has been
conceptualized as relating to numerous cognitive processes includ-
ing attention, memory, perspective taking, theory of mind, abstract
reasoning, cognitive flexibility, generational fluency, and set shift-
ing (Rankin et al., 2005). More concisely, empathy is generally
conceptualized as involving two primary neuropsychological pro-
cesses including the ability to cognitively take the perspective of
others (i.e., cognitive empathy) and the ability to feel the emotions
of others as if they were one’s own (i.e., affective empathy;
Shamay-Tsoory et al., 2009; Shamay-Tsoory, 2011). Studies of
empathy involving persons with brain dysfunction have primarily
focused on the associations between empathy and cognitive flex-
ibility (Grattan et al., 1994; Shamay-Tsoory et al., 2009; Grattan &
Eslinger, 1989), with suggestions that increased cognitive flexibil-
ity allows individuals to take the perspective of others. Other
studies suggest that empathy is related to declarative memory
(Beadle & Tranel, 2013), as well as empathetic concern (Rankin et
al., 2005; Rankin et al., 2006).
Right Hemisphere, Self-Orientation, and Empathy
Empathy has been hypothesized to be related to “self/other
differentiation” and “self/other overlap” (Zaki & Ochsner, 2012).
However, the specific nature of such self/other neuropsychological
processes has not been fully elaborated. Recent research on other
complex human experiences/traits (i.e., transcendence, forgive-
ness) suggests that empathy may have its neuropsychological
foundations in decreased self-orientation associated with the right
hemisphere, and particularly the right parietal lobe (RPL). Re-
search indicates that persons with increasing RPL dysfunction,
from either brain injuries or tumors, report increased spiritual
transcendence (Johnstone, Bodling, Cohen, Christ, & Wegryzn,
2012; Johnstone & Glass, 2008; Urgesi, Aglioti, Skrap, & Fabbro,
2010). This increased transcendence is hypothesized to be related
to decreased self-orientation, or what can be termed increased
“selflessness.” These studies are further supported by single-
photon emission computed tomography (SPECT) research on Bud-
dhist monks and Franciscan nuns who demonstrate decreased RPL
activity during advanced spiritual practices while reporting in-
creased feelings of selflessness (Newberg, Alavi, Baime, Mozley,
& d’Aquili, 1997; Newberg et al., 2001; Newberg, Pourdehand,
Alavi, & d’Aquili, 2003).
Two recently published studies suggest that decreased RPL
functioning is also related to increased willingness to forgive in
populations with both traumatic brain injury (TBI; Johnstone et al.,
2012) and seizures (Johnstone et al., 2014). These studies suggest
that to forgive, one must be willing to decrease focus on the
perceived wrong to the self.
Based on these studies, it is hypothesized that empathy may also
be related to decreased RPL-related self-orientation. Specifically,
it is suggested that one must be selfless to experience spiritual
transcendence, be willing to forgive, and to be empathetic. This
hypothesis is also based on studies that indicate that the right
hemisphere, and particularly the RPL, is primarily related to pro-
cessing information that is related to the self (Austin, 2009;
Northoff et al., 2006). Numerous studies have consistently dem-
onstrated that the right hemisphere is related to several self-
referential processes, such as recognizing pictures of one’s self and
processing autobiographical information/memories (Decety &
Sommerville, 2003; Keenan, McCutcheon, & Pascual-Leone,
2001; Keenan, Nelson, O’Connor, & Pascual-Leone, 2001; Leigh
et al., 2013; Lou et al., 2004; Platek, Wathne, Tierney, & Thom-
son, 2008; Uddin, Molnar-Szakacs, Zaidel, & Iacoboni, 2006).
The relationship between the RPL and self-orientation is also
supported by research with individuals with RPL dysfunction.
They often experience “disorders of the self,” including anosag-
nosia and left-sided neglect, in which they have difficulties
creating/processing a coherent sense of the physical and/or
psychic self (Feinberg & Keenan, 2005b; Feinberg & Keenan,
2005a; McGlynn & Schacter, 1989; Mesulam, 2000). When
considered together, these studies suggest that decreased RPL-
related self-orientation (which can be conceptualized as in-
creased “selflessness”) may also allow individuals to more
easily understand and relate to the experiences of others (i.e., to
be empathetic).
Rationale for Current Study
It was hypothesized that decreased integrity of the RPL (which
is associated with decreased self-orientation) would be related to
increased empathy. To address weaknesses in empathy research to
date, empathy was measured by subjective and objective measures,
and cerebral integrity was measured in terms of both functional
and structural indices. Although the focus of the study was on the
relationship between the RPL and empathy, relationships among
empathy and indices of other cerebral areas were also investigated
to assist in clarifying the neurologic and neuropsychological foun-
dations of empathy.
Methods
Participants
The sample included 31 individuals with TBI referred for out-
patient neuropsychological evaluations at a Midwestern university.
All participants had abnormal clinical radiologic evaluations of the
brain. Neuropsychological tests and self-report measures of empa-
thy were completed by all 31 participants. MRI data for 20
participants was obtained and analyzed for research purposes. Data
were unavailable for the remaining participants due to either
preexisting counterindications (e.g., metal in body; n ⫽ 7) or
technical problems at the time of testing (n ⫽ 4). Chi-square and
t tests conducted between the 20 participants with and 11 partic-
ipants without MRI scanning indicated there were no significant
 SELF-ORIENTATION AND EMPATHY 465

differences in neuropsychological, demographic, or outcome mea- Table 2

sures between the groups, other than marital status (with the group Sample Brain Injury Characteristics
not receiving MRI having a higher percentage of divorce).
The average age of the sample was 34.6 years (SD ⫽ 16.2; range ⫽ N Percent
17– 87). The mean Wechsler Adult Scale of Intelligence III, Full Scale Loss of Consciousness
IQ (WAIS-3 FSIQ) of the sample was in the average range (WAIS-3 Yes 18 58.1
FSIQ ⫽ 93.4; SD ⫽ 15.9; Wechsler, 1997a). The WAIS-3 FSIQ was No 5 16.1
not significantly correlated with any measure of empathy. Demo- Unknown/Missing 8 25.8
Retrograde Amnesia
graphic characteristics of the entire sample are presented in Table 1. Yes 9 29.0
TBI injury severity characteristics are presented in Table 2. No 0 0.0
Unknown/Missing 22 71.0
Posttraumatic Amnesia
Procedure Yes 14 45.2
No 0 0.0
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.

All participants were referred for clinical neuropsychological

Unknown/Missing 17 54.8
This document is copyrighted by the American Psychological Association or one of its allied publishers.

evaluations and had evidence of abnormal neuroradiological eval- Effected Brain Regions on Imaging
uations at the time of their injuries. They were administered Left Frontal 9 29.0
measures of the functional integrity of the brain including intelli- Right Frontal 7 22.6
gence, memory, language, visual-spatial skills, attention, and Left Temporal 7 22.6
Right Temporal 7 22.6
sensory-motor skills by psychometricians (see Table 3). Partici-
Left Parietal 6 19.4
pants provided informed consent and completed subjective mea- Right Parietal 4 12.9
sures of empathy (see Table 4). Participants were then scheduled Occipital 2 6.5
for MRIs (i.e., measures of the structural integrity of the brain),
where they were screened for contraindications to scanning and
also completed an objective measure of empathy. Participants were
compensated $100. The present study was approved by the uni-
versity’s research review board.

Table 1 Measures of Empathy

Sample Demographics
Penner Prosocial Personality Battery (Penner, Fritzsche,
N Percent Craiger, & Freifeld, 1995). This measure was used to assess
several dimensions of empathy and has been used in several
Gender peer-reviewed studies (Finkelstein, Penner, & Brannick, 2005;
Male 21 67.7
Female 10 32.3 Penner, 2002; Penner & Finkelstein, 1998; Skoe, Cumberland,
Handedness Eisenberg, Hansen, & Perry, 2002). The Empathy scale assesses
Right 28 90.3 the ability to feel responsibility for and concern about the welfare
Left 3 9.7 of others. This scale is divided into four different scores, including
Ethnicity
White 28 90.0
a total empathy score, as well as individual scores for each of the
African American 1 3.2 following three dimensions of empathy: Empathetic Concern, Per-
Biracial 2 6.5 spective Taking, and Personal Distress. All questions are rated on
Marital Status
Married 9 29.0
Divorced 5 16.1
Separated 1 3.2 Table 3
Single 16 51.6 Neuropsychological Measure Descriptive Statistics
Education
Some HS 4 12.9 N Min. Max. Mean SD
HS Diploma 13 41.9
Some College 7 22.6 TMT (z score) 31 ⫺19.52 1.18 ⫺3.58 4.86
Bachelor’s Degree 3 9.7 WMS-3 LM I
Master’s Degree 3 9.7 (scaled score) 31 2.00 13.00 7.35 3.01
PhD/JD/MD 1 3.2 BVMT-R Delay
Employment Status (t score) 30 20.00 66.00 46.27 13.77
Currently Employed 12 38.7 JOLO (z score) 31 ⫺1.75 1.08 ⫺0.02 0.86
Student 5 16.1 Letter Fluency
Unemployed, not on disability 12 38.7 (scaled score) 31 3.00 19.00 8.42 3.85
Unemployed, on disability 1 3.2 Agnosia RH
Retired 1 3.2 (errors) 30 0 4 0.70 1.06
Annual Income Agnosia LH
⬍$20,000 25 80.6 (errors) 30 0 2 0.37 0.56
$20,001–$40,000 3 9.7
Note. TMT ⫽ Trail Making Test Part B; WMS-3 LM I ⫽ Wechsler
$40,001–$60,000 1 3.2
Memory Scale III Logical Memory I; BVMT-R ⫽ Benton Visual Memory
$60,001–$80,000 1 3.2
Test Revised; JOLO ⫽ Judgment of Line Orientation; RH ⫽ right hand;
⬎$80,000 1 3.2
LH ⫽ left hand.
 466 JOHNSTONE, COHEN, BRYANT, GLASS, AND CHRIST
Table 4
Subjective and Objective Empathy Measures
Descriptive Statistics
N Min. Max. Mean SD
Prosocial Personality Battery
Total Empathy 31 32 46 39.93 3.79
Perspective Taking 31 10 22 17.32 2.91
Empathic Concern 31 12 19 15.61 2.08
Personal Distress 31 3 12 7.0 2.56
Prisoner’s Dilemma Total Score 20 61 79 71.50 4.35
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.
a five point scale, from 1 (strongly disagree) to 5 (strongly agree).
This document is copyrighted by the American Psychological Association or one of its allied publishers.
The score for each scale is the total number endorsed for each
respective scale. The Total Empathy score is the sum of each of the
following three subscales.
Empathetic concern. This subscale includes four questions
and assesses the degree to which individuals are emotionally
concerned with the well-being of others. It includes questions such
as: “When I see someone being taken advantage of, I feel kind of
protective toward them”; “I am often quite touched by things that
I see happen.” To be consistent with the terminology of previous
studies, Empathetic Concern is conceptualized as affective empa-
thy. Higher scores suggest more empathetic concern. Cronbach’s
alpha for the measure was .67 for the original sample (Penner,
2002; n ⫽ 1,111).
Perspective taking. This subscale includes five questions and
assesses the degree to which individuals can take the perspective
of others. It includes questions such as: “I sometimes try to
understand my friends better by imagining how things look from
their perspective”; “When I am upset at someone, I usually try to
‘put myself in their shoes’ for a while.” To be consistent with the
terminology from previous studies, Perspective Taking is concep-
tualized as cognitive empathy. Higher scores suggest a greater
capacity to take other’s perspective. Cronbach’s alpha for the
measure was .66 for the original sample (Penner, 2002; n ⫽
1,111).
Personal distress. This subscale includes three questions and
assesses the degree to which individuals have difficulties dealing
with stressful situations. Example questions include: “I tend to lose
control during emergencies”; “I am usually pretty effective in
dealing with emergencies”; and “When I see someone who badly
needs help in an emergency, I go to pieces.” Higher scores suggest
a higher degree of distress. Cronbach’s alpha for the measure was
.77 for the original sample (Penner, 2002; n ⫽ 1,111).
Prisoner’s Dilemma task. This computer task that has been
used as an objective measure of empathy/altruism (Batson &
Moran, 1999). Participants are engaged in a task in which they are
told they can cooperate with, or not cooperate with (i.e., defect),
another person (i.e., the same confederate face on the computer
screen for each trial), not knowing if the confederate will them-
selves cooperate or defect. If the participant and confederate both
choose to cooperate, then they both earn $5. If the participant
defects but the confederate cooperates, then the participant earns
$6 and the confederate $1. If the participant cooperates but the
confederate defects, then the participant earns $1 and the confed-
erate $6. The participants are administered 15 trials twice, and the
participant is informed of the confederate’s response after each
trial. The total score is the amount of money earned over all trials.
Higher dollar amounts are indicative of higher levels of coopera-
tion (and theoretically empathy).
Functional and Structural Indices of Cerebral
Integrity
The current study evaluated both the functional and structural
integrity of gross cerebral structures that have been shown to be
related to empathy (i.e., bilateral frontal, temporal, parietal, insular
cortices). Both structural and functional indices of cerebral integ-
rity were used given that neuropsychological tests can provide
different information than neuroradiological techniques (Gomar et
al., 2011; Schmand et al., 2014). For example, cerebral abnormal-
ities were detected in only 18% of MRI scans of persons with TBI,
compared with 40% of SPECT scans and 86% of magnetoen-
cephalography evaluations (Lewine et al., 2007).
Neuropsychological Tests
The following neuropsychological tests were administered as
indices of the general functional integrity of each of the following
cerebral lobes. Specifically, the tests were not used as indicators of
specific cognitive functions, but rather to infer the functional
integrity of the cerebral lobe that is generally associated with the
test (i.e., finger agnosia tests used as indices of parietal lobe
functioning, rather than measures of tactile sensitivity; Lezak,
Howieson, & Loring, 2004). However, it is acknowledged that
only general relationships could be inferred regarding empathy and
neuroanatomical foundations, which was determined to be appro-
priate given the unclear nature of the neurological foundations of
empathy (Zaki & Ochsner, 2012). For all tests, higher scores
indicate more intact ability, other than for the finger agnosia tests
(i.e., higher scores equal worse performance). Age was controlled
for by calculating standard scores for all tests based on age, other
than for finger agnosia.
RPL. The Judgment of Line Orientation (JOLO; Benton,
Hamsher, Varney, & Spreen, 1983) and left-hand finger agnosia
tests were used to infer RPL functioning. The JOLO is a measure
of spatial perception with the total score equaling the number
correct of the 30 total items. For this study, the score is presented
as a z-score calculated according to normative data published in
the test manual, with higher z -scores indicative of better perfor-
mance. The JOLO, and similar measures of the perception of
angulation, have been associated with right hemisphere and par-
ticularly RPL functioning (Benton, Hannay, & Varney, 1975;
Lezak, Howieson, & Loring, 2004; Meador et al., 1993). Left-
sided finger agnosia is a measure of tactile perception that is
associated with the RPL (Lezak et al., 2004). On this test, each of
the five fingers is touched four times and identified by the partic-
ipant, with the score equaling the total number of errors. More
agnosia errors suggest greater RPL dysfunction.
Left parietal lobe. Right-sided finger agnosia is a measure of
tactile perception associated with the left parietal lobe (Lezak et
al., 2004), during which each of the five fingers is touched four
times and identified by the participant, with the score equaling the
total number of errors. More agnosia errors suggest greater left
parietal dysfunction.
Right temporal lobe. The Brief Visuospatial Memory Test
Revised (BVMT-R; Benedict, 1997) is a test of visual memory that
 SELF-ORIENTATION AND EMPATHY 467

has been associated with right temporal lobe functioning (Tranel &
Damasio, 2002). It involves presenting the participant with three
trials during which they are visually exposed to six geometric
figures for 10 s, after which they are asked to draw the figures from
memory. The score is the total number of details recalled for all
three trials, which is presented as a t score based on age-related
normative data (average t score ⫽ 50, SD ⫽ 10).
Left temporal lobe. The Wechsler Memory Scale III
(WMS-3) Logical Memory (LM) I subtest (Wechsler, 1997b) is a
measure of verbal memory that has been associated with left
temporal functioning (Tranel & Damasio, 2002). It involves read-
ing two narrative stories to the participant that they are asked to
remember and repeat immediately after presentation. The score is
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.
(the upsampling is required for use of BrainVoyager’s advanced
segmentation tools). BrainVoyager’s built-in segmentation pro-
cess, which uses local intensity histograms and computed gradient
fields to adaptively calculate WM, GM, and cerebrospinal fluid
(CSF) boundaries, was then applied. Importantly, before proceed-
ing, the resulting segmentations were visually inspected (slice-by-
slice) for accuracy and manually corrected where necessary. Cor-
tical thickness measurements were calculated using a Laplace
method (Jones, Buchbinder, & Aharon, 2000) as implemented
within BrainVoyager. For the current study, cortical volume was
chosen as an index of structural cerebral integrity (i.e., cortical
thickness ⫻ surface area), consistent to other neuroradiological
studies of empathy (Banissy et al., 2012; see Table 5).

the standard score according to age-related norms (i.e., M ⫽ 10,

Output from the previously described segmentation process was
This document is copyrighted by the American Psychological Association or one of its allied publishers.

SD ⫽ 3).
Left frontal lobe. The Delis Kaplan Executive Function Sys- also utilized to create cortical surface reconstructions of each
tem Letter Fluency Test (DKEFS; Delis & Kaplan, 2001) is a hemisphere for each participant. Anatomical alignment of these
measure of verbal fluency that has been associated with left frontal surface representations (and thereby the aforementioned cortical
lobe functioning (Johnstone, Leach, Hickey, Frank, & Rupright, thickness measurements) across participants was accomplished
1995). On this measure, participants state as many words as using BrainVoyager’s automatic cortex-based alignment (CBA)
possible in a 1-min period for three different letters. The total score process (Fischl, Sereno, Tootell, & Dale, 1999; Goebel, Esposito,
is the total number of words generated over the three trials, which & Formisano, 2006). CBA represents an iterative adaptive process
is expressed as a scaled score based on age-related normative data whereby curvature information (representing gyral and sulcal fold-
(M ⫽ 10, SD ⫽ 3). ing patterns) is used to align macroanatomical structures (gyri and
Frontal lobe. The Trail Making Test Part B (Reitan, 1992) sulci) of each participant’s brain to a standard reference brain
has been associated with general frontal lobe functioning (John- provided in BrainVoyager. Following alignment, average cortical
stone et al., 1995). It is a measure of divided attention (also thickness, surface area, and volume measurements were extracted
described as cognitive flexibility) that involves the participant for each major cortical region (frontal, parietal, temporal, insular)
completing a connect-the-dot test, alternating between a series of of each hemisphere (left, right) of each participant.
numbers and letters (i.e., 1 to A, A to 2, 2 to B, B to 3, etc.). The Analyses. Pearson product–moment correlations were con-
result is based on time to complete the measure. For this study, the ducted among the neuropsychological measures and the Penner
score is presented as a z score according to normative data (Hea- Prosocial Personality Battery Empathy scales, and the Prisoner’s
ton, Miller, Taylor, & Grant, 2004), with higher z scores indicative
Dilemma total score (see Table 6). It is noted that age was
of better performance.
accounted for in the analyses through the use of age-corrected
normative data for the neuropsychological measures, other than for
MRI finger agnosia measures. Partial-correlations using age as a cova-
Data acquisition. A 3T Siemens Trio MRI scanner with a riate were conducted among MRI-derived measures of cortical
standard eight-channel head coil was used to obtain high- volume and the Penner scores, and the Prisoner’s Dilemma total
resolution (1 mm3) T1-weighted structural images of the brain. score (see Table 7). Statistical significance was set at 0.05, ac-
Images were collected using a standard T1-weighted pulse se- knowledging weaknesses inherent in using the relatively small
quence (MP-RAGE sequence: repetition time [TR] ⫽ 1,920 ms, sample size.
echo time [TE] ⫽ 4 ms, flip angle ⫽ 8°, number of slices ⫽ 160,
resolution ⫽ 1 mm3).
Data processing and analysis. Data processing and analysis
was carried out using a surface-based approach as implemented
within BrainVoyager QX software (Brain Innovation, Maastricht, Table 5
The Netherlands). (For a more extensive description of the Brain- MRI Cortical Volume Descriptive Statistics
Voyager processing pipeline, see http://support.brainvoyager.com
or Geuze et al., 2008.) First, the structural MRI data for each N Min. Max. Mean SD
participant was rotated into anterior-posterior commissures (AC- Left Hemisphere
PC) coordinates. The skull and dural tissue were then removed by Left Temporal 20 30,628.05 53,710.81 42,241.44 5,425.12
manually deleting voxels containing non-neural tissue. Next, a Left Frontal 20 41,317.70 79,798.30 59,528.84 8,539.29
Sigma filter and BrainVoyager’s automatic intensity inhomogene- Left Parietal 20 23,028.63 37,681.60 31,215.89 3,722.84
Left Insula 20 6,319.09 9,724.08 8,121.69 900.11
ity correction tool (IIHC) were applied to enhance grey/white Right Hemisphere
matter (GM/WM) tissue contrast and correct for spatial intensity Right Temporal 19 30,052.48 47,168.65 40,602.12 4,330.72
inhomogeneities, respectively. Noncortical structures (e.g., ventri- Right Parietal 20 44,251.37 81,739.61 62,146.05 8,641.45
cles, subcortical nuclei) were then removed, and the resulting Right Frontal 20 23,802.39 34,333.18 28,024.57 3,026.48
Right Insula 20 8,123.95 11,413.30 9,915.95 894.49
image was upsampled from 1 to 0.5 mm3 using sinc interpolation
 468 JOHNSTONE, COHEN, BRYANT, GLASS, AND CHRIST

Table 6 related to increased rather than decreased RPL functional integ-

Correlation Analyses Among Neuropsychological Indices and rity. The process of being empathetic apparently is not associated
Empathy Measures with a decreased focus on the self, which allows for an increased
focus on others. Instead, it appears that to empathize with others
Prisoner’s that it is necessary to have an intact and relatively strong sense of
Neuropsychological Total EC PT PD Dilemma
the self. That is, if one does not have an intact sense of the self,
Left Temporal then one cannot fully understand and emotionally relate to the
WMS-3 LM I 0.24 ⫺0.16 0.36ⴱ 0.14 0.28 experiences of others. As stated by Cozolino:
Left Frontal
DKEFS Letter Fluency 0.52ⴱ 0.42ⴱ 0.42ⴱ ⫺0.08 0.04 empathy is an attempt to experience the inner life of another while
Left Parietal retaining objectivity. In other words, we hold our own perspective in
Right Finger Agnosia 0.07 0.11 ⫺0.06 0.08 ⫺0.30
mind while simultaneously imagining what it is like to be the other. In
Right Temporal
BVMT-R total 0.03 ⫺0.24 0.09 0.12 0.07 order to have empathy, we need to maintain an awareness of our inner
Right Parietal world as we imagine the inner world of others. (Cozolino, 2006, p.
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.

JOLO 0.04 0.38ⴱ 0.12 ⫺0.49ⴱ 0.20 203)

This document is copyrighted by the American Psychological Association or one of its allied publishers.

Left Finger Agnosia ⫺0.21 ⫺0.37ⴱ ⫺0.27 0.37ⴱ 0.15

Frontal Although the results were contrary to those hypothesized, they
Trail Making Test 0.35ⴱ ⫺0.01 0.39ⴱ 0.31 0.16 support others studies that suggest that relatively stronger right
Note. Total ⫽ Penner Prosocial Personality Total score; EC ⫽ Empa- hemisphere functioning, and particularly RPL functioning, is re-
thetic Concern; PT ⫽ Perspective Taking; PD ⫽ Personal Distress; Pris- lated to increased empathy (Decety & Batson, 2009; Grattan &
oner’s Dilemma ⫽ Prisoner’s Dilemma Total Earnings; WMS-3 LM I ⫽ Eslinger, 1989; Shamay-Tsoory et al., 2004; Shamay-Tsoory et al.,
Wechsler Memory Scale III Logical Memory I; BVMT-R ⫽ Benton Visual 2009). For example, Shamay-Tsoory and colleagues (2004) re-
Memory Test Revised; DKEFS ⫽ Delis Kaplan Executive Function
Scales; JOLO ⫽ Judgment of Line Orientation. ported that individuals with RPL lesions, but not left parietal lobe
ⴱ lesions, demonstrated significantly decreased empathy. Similarly,
p ⬍ .05.
Grattan and Eslinger (1989) found that the JOLO (i.e., the same
measure used in the current study to infer RPL dysfunction) was
significantly and positively related to empathy in a sample of
Results individuals with brain dysfunction (r ⫽ .30 vs. r ⫽ .38 in the
current study). However, they conceptualized this test as a measure
Neuropsychological Indices of perceptual judgment and did not mention this finding in the
Contrary to hypotheses, the results suggest that empathy is abstract given the focus of their study was on the frontal lobes. In
generally related to increased functional integrity of the RPL. contrast, in the current study, the JOLO was used to infer the
Specifically, measures of increased RPL functioning (i.e., JOLO functional integrity of the RPL. Whereas Grattan and Eslinger
and left-hand finger agnosia tests) were significantly correlated concluded that “impairments in visual-spatial perception could
with both increased affective empathy and less personal distress. potentially be a shared variable among the flexibility and empathy
Measures associated with frontal lobe functioning, and particu- measures,” it is suggested that it may be more appropriate to
larly the left frontal lobe (i.e., Trail Making Test and DKEFS interpret these tests as indicators of general RPL functional integ-
Letter Fluency subtest), were significantly and positively corre- rity. Based on this premise, it can then be inferred that empathy is
lated with cognitive empathy. significantly related to two measures of RPL integrity (i.e., JOLO,
A measure of left temporal lobe functioning (i.e., WMS-3 LM I left finger agnosia), rather than suggesting that empathy is related
scale) was significantly and positively associated with cognitive
empathy.
Table 7
No neuropsychological measures were related to Prisoner’s Di-
Correlation Analysis Among MRI Cortical Volume and
lemma scores.
Empathy Measures
MRI Indices Prisoner’s
MRI Brain Region Total EC PT PD Dilemma
Right and left insular cortices were the only cerebral structures
shown by MRI to be significantly related to subjective measures of Left Hemisphere
empathy (i.e., Total Empathy, Personal Distress). No MRI indices Left Temporal 0.32 0.18 0.19 0.05 0.04
Left Frontal 0.12 0.08 ⫺0.13 0.20 ⫺0.11
of cerebral integrity were significantly related to Prisoner’s Di- Left Parietal 0.33 0.27 0.25 ⫺0.04 0.04
lemma scores. Left Insula 0.48ⴱ 0.06 0.25 0.46ⴱ 0.15
Right Hemisphere
Right Temporal 0.12 ⫺0.10 ⫺0.13 0.26 ⫺0.07
Discussion Right Frontal ⫺0.15 ⫺0.17 ⫺0.23 0.03 ⫺0.12
Right Parietal ⫺0.02 ⫺0.04 ⫺0.02 ⫺0.07 0.12
Empathy and the RPL Right Insula 0.43ⴱ ⫺0.18 0.29 0.56ⴱ 0.04
Note. Total ⫽ Total Empathy; EC ⫽ Empathetic Concern; PT ⫽ Per-
The primary goal of the study was to determine if decreased spective Taking; PD ⫽ Personal Distress; Prisoner’s Dilemma ⫽ Prison-
RPL functioning was related to increased empathy. Contrary to the er’s Dilemma Total Earnings.
ⴱ
hypothesis, the results suggest that certain aspects of empathy are p ⬍ .05.
 SELF-ORIENTATION AND EMPATHY
to perceptual judgment and left-sided tactile sensitivity. If concep-
tualized in this manner, it can then be inferred that other RPL
functions, such as general self-orientation processes (Austin, 2009;
Decety & Sommerville, 2003; Leigh et al., 2013; Lou et al., 2004;
Uddin et al., 2006; Feinberg & Keenan, 2005b, 2005a; McGlynn &
Schacter, 1989; Mesulam, 2000) are also likely to be diminished.
Of note, the results also indicated that decreased RPL function-
ing (i.e., JOLO, left-sided finger agnosia) is associated with higher
levels of personal distress. That is, less intact RPL functioning
(and inferentially the sense of self) is associated with a decreased
ability to keep composure under stressful situations (e.g., “I tend to
lose control during emergencies”; “When I see someone who
badly needs help in an emergency, I go to pieces”). This suggests
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.
that a stronger sense of the self allows individuals to retain com-
This document is copyrighted by the American Psychological Association or one of its allied publishers.
posure in stressful situations and increase their ability to focus on
others. This finding is consistent with beliefs that one must have a
strong sense of the self to empathize with others (Cozolino, 2006).
Empathy and Other Cerebral
Structures/Neuropsychological Functions
In the current study, empathy (i.e., affective, cognitive, total)
was associated with frontal lobe functioning (and particularly the
left frontal lobe), consistent with several other studies (Decety &
Batson, 2009; Decety & Jackson, 2004; Grattan et al., 1994;
Grattan & Eslinger, 1989; Grattan & Eslinger, 1992; Rankin et al.,
2005; Rankin et al., 2006; Shamay-Tsoory et al., 2004; Shamay-
Tsoory et al., 2009). In general, current and previous results
consistently suggest that cognitive flexibility associated with fron-
tal lobe functioning is necessary for individuals to take the cogni-
tive perspective of others.
The results also support previous studies that indicate that the
temporal lobes are involved in empathy (Decety & Batson, 2009;
Leigh et al., 2013; Rankin et al., 2006). It has been suggested that
empathy may be related to the emotional processing that has been
shown to be related to both the temporal lobes and limbic system,
given their close neural connectivity (Lee & Siegle, 2012; Lezak et
al., 2004; Rankin et al., 2005). However, somewhat unexpectedly,
in the current study, cognitive empathy was associated with in-
creased left temporal lobe functioning, whereas affective empathy
was not. If anything, it could be hypothesized that affective em-
pathy would be related to temporal lobe functioning. The rationale
for this finding is unclear.
Empathy and Structural Indices of Cerebral Integrity
The results indicate that the bilateral insular cortices were the
only structural indices that were significantly associated with
empathy (i.e., Total Empathy and Personal Distress). This finding
is generally consistent with several other studies that implicate the
insula in the experience of empathy (Banissy et al., 2012;
Mutschler et al., 2013). It is speculated that the insula may be
related to empathy given that it has been associated with emotional
processing (and particularly negative emotions such as personal
distress), awareness of internal bodily states (i.e., self-orientation
of internal states), and connecting physiological states of the body
with social emotions such as empathy (Craig, 2009; Critchley,
Wiens, Rotshtein, Ohman, & Dolan, 2004; Karnath, Baier, &
Nägele, 2005).
469
The lack of significant findings between empathy and other
cerebral areas does not suggest that empathy is solely based in the
insula. The lack of findings is more likely related to the fact that
many individuals with even severe TBI have normal neuroradio-
logical findings (Brasure et al., 2012; Lobato et al., 1986). This
suggests that quantitative structural MRI data may help differen-
tiate groups with brain dysfunction for comparison (e.g., frontal vs.
nonfrontal lesions; Grattan, Bloomer, Archambault, & Eslinger
1994), but may not be appropriate for correlational analyses such
as those conducted in the current study. Regardless, the current
results clearly suggest that future empathy research will benefit
from focusing on the role of the insula.
Self-Report Versus Objective Measures of Empathy
The results indicate that self-report measures of empathy (i.e.,
Penner’s Prosocial Personality measure) were significantly related
to neuropsychological indices of cerebral integrity, although the
objective measure (i.e., Prisoner’s Dilemma) was not. This finding
may be reflective of the “artificiality” associated with using ex-
perimental tasks to measure empathy (Zaki & Ochsner, 2012), and
particularly for persons with TBI. Although subjective and objec-
tive measures of a construct may frequently be administered to-
gether to assure the construct validity of an attribute (e.g., empa-
thy), in this case the constructs measured by the Penner Prosocial
Personality Measure and Prisoner’s Dilemma appear to be differ-
ent.
Limitations and Future Directions
It is acknowledged that empathetic processes are much more
complicated from neuroanatomical and neuropsychological per-
spectives than the conclusions presented in this article. However,
given the weaknesses in empathy research to date (Zaki &
Ochsner, 2012), the current results can provide suggestions for
future research, and particularly how self-orientation relates to
empathy. The current study suggests that different neuropsycho-
logical processes associated with different neurological networks
interact in the experience of empathy (e.g., RPL-related self-
orientation; frontal lobe-related cognitive flexibility; temporal
lobe-related emotional processing; insula-related processing of
internal physical states and social emotions).
To date researchers have generally conceptualized empathy as
involving processes such as “self/other discrimination,” “self-
reflection,” “self/other overlap,” and “theory of mind” (Shamay-
Tsoory et al., 2009; Zaki & Ochsner, 2012). However, these terms
are relatively nonspecific and such terminology may not best
explain the manner by which individuals incorporate the experi-
ences of others into the experience of the self. A more nuanced
explanation for the manner by which specific dimensions of self-
orientation relate to empathy is needed. Just as “language” can be
used as a general umbrella term for other more specific commu-
nication skills (e.g., expression, comprehension, semantics, syntax,
etc.), “self-orientation” may be best conceptualized as a general
umbrella descriptor for more specific self-oriented processes (e.g.,
autobiographical memory, body schema perception, self-
awareness, self-recognition, self-evaluation, etc.). In fact, a general
model of self-orientation has been proposed, with suggestions that
the self can be conceptualized in terms of a physical self, mental
self, and autobiographical self (Northoff et al., 2006).
 470 JOHNSTONE, COHEN, BRYANT, GLASS, AND CHRIST
Further research regarding the expression of increased and de-
creased self-orientation processes is also warranted given the mul-
tiple studies that suggest that decreased RPL functioning (as mea-
sured by the JOLO and/or finger agnosia tests) is associated with
increased transcendence (Johnstone et al., 2012; Johnstone &
Glass, 2008) and increased willingness to forgive (Johnstone et al.,
2012; Johnstone et al., 2014), but decreased empathy (Grattan et
al., 1994; current study). Future research may benefit from deter-
mining how increased and decreased self-orientation may be re-
lated to other human experiences (e.g., altruism, antisocial person-
ality, etc.), as well as the manner in which self- and other-oriented
neuropsychological processes interact in different human experi-
ences/relationships (Austin, 2009; Fox et al., 2005; Ochsner et al.,
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.
2004).
This document is copyrighted by the American Psychological Association or one of its allied publishers.
A primary limitation of the current study is the relatively small
sample size, as the use of a larger sample may lead to a more clear
understanding of the specific nature of empathy (e.g., allow for
investigation of more specific brain regions). The results are fur-
ther limited in that the current sample included persons with TBI,
which is an injury that typically involves generalized impairment
throughout the brain (Lezak et al., 2004). As a result, only general
inferences can be drawn regarding the neuroanatomical and neu-
ropsychological bases of empathy. Relatedly, the current study is
limited as the MRI data involving cortical volume was viewed by
examining large regions as a whole (i.e., left frontal lobe) rather
than examining more specific regions within each brain region that
have been suggested to be implicated in different dimensions of
empathy (Eslinger, 1998; Shamay-Tsoory et al., 2009). Not focus-
ing on more specific brain regions may have decreased the strength
of the observed relationships and may have contributed to the
insignificant MRI findings.
References
Austin, J. H. (2009). Selfless insight. Cambridge, MA: MIT Press.
Banissy, M. J., Kanai, R., Walsh, V., & Rees, G. (2012). Inter-individual
differences in empathy are reflected in human brain structure. Neuro-
Image, 62, 2034 –2039. http://dx.doi.org/10.1016/j.neuroimage.2012.05
.081
Batson, C. D., & Moran, T. (1999). Empathy induced altruism in a
prisoner’s dilemma. European Journal of Social Psychology, 29, 909 –
924. http://dx.doi.org/10.1002/(SICI)1099-0992(199911)29:7⬍909::
AID-EJSP965⬎3.0.CO;2-L
Beadle, J. N., & Tranel, D. (2013). Empathy in hippocampal amnesia.
Frontiers in Psychology, 4, 091.
Benedict, R. (1997). Brief Visuospatial Memory Test-Revised professional
manual. Odessa, FL: Psychological Assessment Resources.
Benton, A. L., Hamsher, K. D., Varney, N. R., & Spreen, O. (1983).
Judgement of line orientation manual. New York, NY: Oxford Univer-
sity Press.
Benton, A., Hannay, H. J., & Varney, N. R. (1975). Visual perception of
line direction in patients with unilateral brain disease. Neurology, 25,
907–910. http://dx.doi.org/10.1212/WNL.25.10.907
Brasure, M., Lamberty, G. J., Sayer, N. A., Nelson, N. W., MacDonald, R.,
Ouellette, J., . . . Wilt, T. J. (2012). Multidisciplinary postacute rehabil-
itation for moderate to severe traumatic brain injury in adults. Agency for
Healthcare Research and Quality (AHRQ) Comparative Effectiveness
Reviews, 72, ES1–ES20.
Cozolino, L. (2006). The neuroscience of human relationships. New York,
NY: Norton.
Craig, A. D. (2009). How do you feel—Now? The anterior insula and
human awareness. Nature Reviews Neuroscience, 10, 59 –70. http://dx
.doi.org/10.1038/nrn2555
Critchley, H. D., Wiens, S., Rotshtein, P., Ohman, A., & Dolan, R. J.
(2004). Neural systems supporting interoceptive awareness. Nature Neu-
roscience, 7, 189 –195. http://dx.doi.org/10.1038/nn1176
Decety, J., & Batson, C. D. (2009). Empathy and morality: Integrating
social and neuroscience approaches. In J. Braeckman, J. Verplaetse, & J.
De Schrijver (Eds.), The moral brain (pp. 109 –127). Berlin: Springer-
Verlag. http://dx.doi.org/10.1007/978-1-4020-6287-2_5
Decety, J., & Jackson, P. L. (2004). The functional architecture of lesion
empathy. Behavior, Cognition, & Neuroscience, 3, 71–100.
Decety, J., & Jackson, P. L. (2006). A social-neuroscience perspective on
empathy. Current Directions in Psychological Science, 15, 54 –58.
http://dx.doi.org/10.1111/j.0963-7214.2006.00406.x
Decety, J., & Sommerville, J. A. (2003). Shared representations between
self and other: A social cognitive neuroscience view. Trends in Cogni-
tive Sciences, 7, 527–533. http://dx.doi.org/10.1016/j.tics.2003.10.004
Delis, D., & Kaplan, E. (2001). Delis Kaplan Executive Function System.
San Antonio, TX: The Psychological Corporation.
de Sousa, A., McDonald, S., Rushby, J., Li, S., Dimoska, A., & James, C.
(2010). Why don’t you feel how I feel? Insight into the absence of
empathy after severe traumatic brain injury. Neuropsychologia, 48,
3585–3595. http://dx.doi.org/10.1016/j.neuropsychologia.2010.08.008
Eslinger, P. J. (1998). Neurological and neuropsychological bases of em-
pathy. European Neurology, 39, 193–199. http://dx.doi.org/10.1159/
000007933
Feinberg, T. E., & Keenan, J. P. (2005a). The lost self: Pathologies of the
brain and identity. New York, NY: Oxford University Press. http://dx
.doi.org/10.1093/acprof:oso/9780195173413.001.0001
Feinberg, T. E., & Keenan, J. P. (2005b). Where in the brain is the self?
Consciousness and Cognition: An International Journal, 14, 661– 678.
http://dx.doi.org/10.1016/j.concog.2005.01.002
Finkelstein, M. A., Penner, L. A., & Brannick, M. T. (2005). Motive, role
identity, and prosocial personality as predictors of volunteer activity.
Social Behavior and Personality, 33, 403– 418. http://dx.doi.org/
10.2224/sbp.2005.33.4.403
Fischl, B., Sereno, M. I., Tootell, R. B., & Dale, A. M. (1999). High-
resolution intersubject averaging and a coordinate system for the cortical
surface. Human Brain Mapping, 8, 272–284. http://dx.doi.org/10.1002/
(SICI)1097-0193(1999)8:4⬍272::AID-HBM10⬎3.0.CO;2-4
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 Na-
tional Academy of Sciences of the United States of America, 102,
9673–9678. http://dx.doi.org/10.1073/pnas.0504136102
Geuze, E., Westenberg, H. G. M., Heinecke, A., de Kloet, C. S., Goebel,
R., & Vermetten, E. (2008). Thinner prefrontal cortex in veterans with
posttraumatic stress disorder. NeuroImage, 41, 675– 681. http://dx.doi
.org/10.1016/j.neuroimage.2008.03.007
Goebel, R., Esposito, F., & Formisano, E. (2006). Analysis of functional
image analysis contest (FIAC) data with BrainVoyager QX: From
single-subject to cortically aligned group general linear model analysis
and self-organizing group independent component analysis. Human
Brain Mapping, 27, 392– 401. http://dx.doi.org/10.1002/hbm.20249
Gomar, J. J., Bobes-Bascaran, M. T., Conejero-Goldberg, C., Davies, P.,
Goldberg, T. E., & the Alzheimer’s Disease Neuroimaging Initiative.
(2011). Utility of combinations of biomarkers, cognitive markers, and
risk factors to predict conversion from mild cognitive impairment to
Alzheimer disease in patients in the Alzheimer’s disease neuroimaging
initiative. Archives of General Psychiatry, 68, 961–969. http://dx.doi
.org/10.1001/archgenpsychiatry.2011.96
Grattan, L. M., Bloomer, R. H., Archambault, F. X., & Eslinger, P. J.
(1994). Cognitive flexibility and empathy after frontal lobe lesions.
 SELF-ORIENTATION AND EMPATHY
Neuropsychiatry, Neuropsychology, & Behavioral Neurology, 7, 251–
259.
Grattan, L. M., & Eslinger, P. J. (1989). Higher cognition and social
behavior: Changes in cognitive flexibility and empathy after cerebral
lesions. Neuropsychology, 3, 175–185.
Grattan, L. M., & Eslinger, P. J. (1992). Long-term psychological conse-
quences of childhood frontal lobe lesion in patient DT. Brain and
Cognition, 20, 185–195. http://dx.doi.org/10.1016/0278-2626
(92)90068-W
Heaton, R. K., Miller, W., Taylor, M. J., & Grant, I. (2004). Revised
comprehensive norms for an expanded Halstead-Reitan Battery. Lutz,
FL: Psychological Assessment Resources.
Johnstone, B., Bayan, S., Gutierrez, L., Lardizabal, D., Lanigar, S., Yoon,
D. P., & Judd, K. (2014). Neuropsychological correlates of forgiveness.
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.
Religion, Brain & Behavior. http://dx.doi.org/10.1080/2153599X.2014
This document is copyrighted by the American Psychological Association or one of its allied publishers.
.880069
Johnstone, B., Bodling, A., Cohen, D., Christ, S. E., & Wegrzyn, A.
(2012). Right parietal lobe-related “selflessness” as the neuropsycholog-
ical basis of spiritual transcendence. International Journal for the Psy-
chology of Religion, 22, 267–284. http://dx.doi.org/10.1080/10508619
.2012.657524
Johnstone, B., & Glass, B. A. (2008). Evaluation of a neuropsychological
model of spirituality in persons with traumatic brain injury. Zygon, 43,
861– 874. http://dx.doi.org/10.1111/j.1467-9744.2008.00964.x
Johnstone, B., Leach, L. R., Hickey, M. L., Frank, R. G., & Rupright, J.
(1995). Some objective measurements of frontal lobe deficits following
traumatic brain injury. Applied Neuropsychology, 2, 24 –28. http://dx.doi
.org/10.1207/s15324826an0201_4
Jones, S. E., Buchbinder, B. R., & Aharon, I. (2000). Three-dimensional
mapping of cortical thickness using Laplace’s equation. Human Brain
Mapping, 11, 12–32. http://dx.doi.org/10.1002/1097-0193(200009)11:
1⬍12::AID-HBM20⬎3.0.CO;2-K
Karnath, H. O., Baier, B., & Nägele, T. (2005). Awareness of the functioning
of one’s own limbs mediated by the insular cortex? The Journal of Neuro-
science, 25, 7134 –7138. http://dx.doi.org/10.1523/JNEUROSCI
.1590-05.2005
Keenan, J. P., McCutcheon, N. P., & Pascual-Leone, A. (2001). Functional
magnetic resonance imaging and event-related potentials suggest right
prefrontal activation for self-related processing. Brain and Cognition,
47, 87–91.
Keenan, J. P., Nelson, A., O’Connor, M., & Pascual-Leone, A. (2001).
Self-recognition and the right hemisphere. Nature, 409, 305. http://dx
.doi.org/10.1038/35053167
Lee, K. H., & Siegle, G. J. (2012). Common and distinct brain networks
underlying explicit emotional evaluation: A meta-analytic study. Social
Cognitive and Affective Neuroscience, 7, 521–534. http://dx.doi.org/
10.1093/scan/nsp001
Leigh, R., Oishi, K., Hsu, J., Lindquist, M., Gottesman, R. F., Jarso, S., . . .
Hillis, A. E. (2013). Acute lesions that impair affective empathy. Brain:
A Journal of Neurology, 136, 2539 –2549. http://dx.doi.org/10.1093/
brain/awt177
Lewine, J. D., Davis, J. T., Bigler, E. D., Thoma, R., Hill, D., Funke, M.,
. . . Orrison, W. W. (2007). Objective documentation of traumatic brain
injury subsequent to mild head trauma: Multimodal brain imaging with
MEG, SPECT, and MRI. The Journal of Head Trauma Rehabilitation,
22, 141–155. http://dx.doi.org/10.1097/01.HTR.0000271115.29954.27
Lezak, M. D., Howieson, D. B., & Loring, D. W. (2004). Neuropsycho-
logical assessment (4th ed.). New York, NY: Oxford Press.
Lobato, R. D., Sarabia, R., Rivas, J. J., Cordobes, F., Castro, S., Muñoz,
M. J., . . . Lamas, E. (1986). Normal computerized tomography scans in
severe headinjury. Prognostic and clinical management implications.
Journal of Neurosurgery, 65, 784 –789. http://dx.doi.org/10.3171/jns
.1986.65.6.0784
471
Logothetis, N. K. (2008). What we can do and what we cannot do with
fMRI. Nature, 453, 869 – 878. http://dx.doi.org/10.1038/nature06976
Lou, H. C., Luber, B., Crupain, M., Keenan, J. P., Nowak, M., Kjaer,
T. W., . . . Lisanby, S. H. (2004). Parietal cortex and representation of
the mental Self. Proceedings of the National Academy of Sciences of the
United States of America, 101, 6827– 6832. http://dx.doi.org/10.1073/
pnas.0400049101
McGlynn, S. M., & Schacter, D. L. (1989). Unawareness of deficits in
neuropsychological syndromes. Journal of Clinical and Experimental
Neuropsychology, 11, 143–205. http://dx.doi.org/10.1080/
01688638908400882
Meador, K. J., Moore, E. E., Nichols, M. E., Abney, O. L., Taylor, H. S.,
Zamrini, E. Y., & Loring, D. W. (1993). The role of cholinergic systems
in visuospatial processing and memory. Journal of Clinical and Exper-
imental Neuropsychology, 15, 832– 842.
Mesulam, M.-M. (2000). Behavioral neuroanatomy. In M.-M. Mesulam
(Ed.), Principles of behavioral and cognitive neurology (2nd ed.). New
York, NY: Oxford Press.
Mutschler, I., Reinbold, C., Wankerl, J., Seifritz, E., & Ball, T. (2013).
Structural basis of empathy and the domain general region in the anterior
insular cortex. Frontiers in Human Neuroscience, 7, 177. http://dx.doi
.org/10.3389/fnhum.2013.00177
Newberg, A. B., Alavi, A., Baime, M., Mozley, P. D., & d’Aquili, E. G.
(1997). The measurement of cerebral blood flow during the complex
cognitive task of mediation using HMPAO-SPECT imaging. Journal of
Nuclear Medicine, 38, 95.
Newberg, A., Alavi, A., Baime, M., Pourdehnad, M., Santanna, J., &
d’Aquili, E. (2001). The measurement of regional cerebral blood flow
during the complex cognitive task of meditation: A preliminary SPECT
study. Psychiatry Research, 106, 113–122. http://dx.doi.org/10.1016/
S0925-4927(01)00074-9
Newberg, A., Pourdehnad, M., Alavi, A., & d’Aquili, E. G. (2003).
Cerebral blood flow during meditative prayer: Preliminary findings and
methodological issues. Perceptual and Motor Skills, 97, 625– 630. http://
dx.doi.org/10.2466/pms.2003.97.2.625
Northoff, G., Heinzel, A., de Greck, M., Bermpohl, F., Dobrowolny, H., &
Panksepp, J. (2006). Self-referential processing in our brain—A meta-
analysis of imaging studies on the self. NeuroImage, 31, 440 – 457.
http://dx.doi.org/10.1016/j.neuroimage.2005.12.002
Ochsner, K. N., Knierim, K., Ludlow, D. H., Hanelin, J., Ramachandran,
T., Glover, G., & Mackey, S. C. (2004). Reflecting upon feelings: An
fMRI study of neural systems supporting the attribution of emotion to
self and other. Journal of Cognitive Neuroscience, 16, 1746 –1772.
http://dx.doi.org/10.1162/0898929042947829
Penner, L. A. (2002). Dispositional and organizational influences on sus-
tained volunteerism: An interactionist perspective. Journal of Social
Issues, 58, 447– 467. http://dx.doi.org/10.1111/1540-4560.00270
Penner, L. A., & Finkelstein, M. A. (1998). Dispositional and structural
determinants of volunteerism. Journal of Personality and Social Psy-
chology, 74, 525–537. http://dx.doi.org/10.1037/0022-3514.74.2.525
Penner, L. A., Fritzsche, B. A., Craiger, J. P., & Freifeld, T. S. (1995).
Measuring the prosocial personality. In J. Butcher & C. D. Spielberger
(Eds.), Advances in personality assessment (Vol. 10). Hillsdale, NJ:
LEA.
Platek, S. M., Wathne, K., Tierney, N. G., & Thomson, J. W. (2008).
Neural correlates of self-face recognition: An effect-location meta-
analysis. Brain Research, 1232, 173–184. http://dx.doi.org/10.1016/j
.brainres.2008.07.010
Preston, S. D., & de Waal, F. B. (2002). Empathy: Its ultimate and
proximate bases. Behavioral and Brain Sciences, 25, 1–20.
Rankin, K. P., Gorno-Tempini, M. L., Allison, S. C., Stanley, C. M., Glenn,
S., Weiner, M. W., & Miller, B. L. (2006). Structural anatomy of
empathy in neurodegenerative disease. Brain: A Journal of Neurology,
129, 2945–2956. http://dx.doi.org/10.1093/brain/awl254
 472 JOHNSTONE, COHEN, BRYANT, GLASS, AND CHRIST
Rankin, K. P., Kramer, J. H., & Miller, B. L. (2005). Patterns of cognitive
and emotional empathy in frontotemporal lobar degeneration. Cognitive
and Behavioral Neurology, 18, 28 –36. http://dx.doi.org/10.1097/01.wnn
.0000152225.05377.ab
Reitan, R. M. (1992). Trail making test: Manual for administration and
scoring. Tucson, AZ: Reitan Neuropsychological Laboratory.
Rorden, C., & Karnath, H.-O. (2004). Using human brain lesions to infer
function: A relic from a past era in the fMRI age? Nature Reviews
Neuroscience, 5, 812– 819. http://dx.doi.org/10.1038/nrn1521
Schmand, B., Rienstra, A., Tamminga, H., Richard, E., van Gool, W. A.,
Caan, M. W., & Majoie, C. B. (2014). Responsiveness of magnetic
resonance imaging and neuropsychological assessment in memory clinic
patients. Journal of Alzheimer’s Disease, 40, 409 – 418.
Shamay-Tsoory, S. G. (2011). The neural bases for empathy. The Neuro-
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.
scientist, 17, 18 –24. http://dx.doi.org/10.1177/1073858410379268
This document is copyrighted by the American Psychological Association or one of its allied publishers.
Shamay-Tsoory, S. G., Aharon-Peretz, J., & Perry, D. (2009). Two systems
for empathy: A double dissociation between emotional and cognitive
empathy in inferior frontal gyrus versus ventromedial prefrontal lesions.
Brain: A Journal of Neurology, 132, 617– 627. http://dx.doi.org/
10.1093/brain/awn279
Shamay-Tsoory, S. G., Tomer, R., Goldsher, D., Berger, B. D., & Aharon-
Peretz, J. (2004). Impairment in cognitive and affective empathy in
patients with brain lesions: Anatomical and cognitive correlates. Journal
of Clinical and Experimental Neuropsychology, 26, 1113–1127. http://
dx.doi.org/10.1080/13803390490515531
Skoe, E. A., Cumberland, A., Eisenberg, N., Hansen, K., & Perry, J.
(2002). The influences of sex and gender-role identity on moral cogni-
tion and prosocial personality traits. Sex Roles, 46, 295–309. http://dx
.doi.org/10.1023/A:1020224512888
Tranel, D., & Damasio, A. R. (2002). Neurobiological foundations of
human memory. In A. D. Baddeley, M. D. Kopelman, & B. A. Wilson
(Eds.), The handbook of memory disorders (2nd ed., pp. 17–56). Chich-
ester: Wiley.
Uddin, L. Q., Molnar-Szakacs, I., Zaidel, E., & Iacoboni, M. (2006). rTMS
to the right inferior parietal lobule disrupts self-other discrimination.
Social Cognitive and Affective Neuroscience, 1, 65–71. http://dx.doi.org/
10.1093/scan/nsl003
Urgesi, C., Aglioti, S. M., Skrap, M., & Fabbro, F. (2010). The spiritual
brain: Selective cortical lesions modulate human self-transcendence.
Neuron, 65, 309 –319. http://dx.doi.org/10.1016/j.neuron.2010.01.026
Wechsler, D. (1997a). WAIS III. Administration and scoring manual. San
Antonio, TX: The Psychological Corporation.
Wechsler, D. (1997b). WMS-III. Administration and scoring manual. San
Antonio, TX: The Psychological Corporation.
Zaki, J., & Ochsner, K. N. (2012). The neuroscience of empathy: Progress,
pitfalls and promise. Nature Neuroscience, 15, 675– 680. http://dx.doi
.org/10.1038/nn.3085
Received March 7, 2014
Revision received July 24, 2014
Accepted September 22, 2014 䡲