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6 (4) p.327-335
Mirror neuron system involvement in empathy: A critical look at the evidence
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Mirror neuron system involvement in empathy: A
critical look at the evidence
Amee D. Baird a , Ingrid E. Scheffer b c & Sarah J. Wilson a c
a Psychological Sciences, University of Melbourne, Melbourne, Australia
b Department of Medicine, University of Melbourne, Melbourne, Australia
c Epilepsy Research Centre, Austin Health, Melbourne, Australia
Available online: 19 Sep 2011
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SOCIAL NEUROSCIENCE, 2011, 6 (4), 327–335
© 2011 Psychology Press, an imprint of the Taylor & Francis Group, an Informa business
www.psypress.com/socialneuroscience DOI: 10.1080/17470919.2010.547085
PSNS Mirror neuron system involvement in empathy:
A critical look at the evidence
Mirror Neuron System In Empathy
Amee D. Baird1, Ingrid E. Scheffer2,3, and Sarah J. Wilson1,3
1Psychological Sciences, University of Melbourne, Melbourne, Australia
2Department of Medicine, University of Melbourne, Melbourne, Australia
3Epilepsy Research Centre, Austin Health, Melbourne, Australia
It has been proposed that the human mirror neuron system (MNS) plays an integral role in mediating empathy. In
this review, we critically examine evidence from three bodies of research that have been cited as supporting this
notion: (1) behavioral studies that have examined the relationship between imitation and empathy, (2) findings
from functional neuroimaging studies that report a positive correlation between MNS activation and self-report on
an empathy questionnaire, and (3) observations of impaired imitation and empathy in autism spectrum disorders
(ASD). In addition, we briefly review lesion studies of the neural correlates of imitation and empathy. Current
evidence suggests that the MNS is broadly involved in empathy, but at this stage there has been limited consider-
ation of its various forms, including motor, emotional, and cognitive empathy. There are also various forms of
imitation, encompassing emotional and non-emotional, automatic, and voluntary actions. We propose that the
relationship between imitation and empathy may vary depending on the specific type of each of these abilities.
Furthermore, these abilities may be mediated by partially distinct neural networks, which involve the MNS to a
Keywords: Empathy; Mirror neuron system; Imitation; Autism spectrum disorders (ASD).
The initial discovery of the human mirror neuron sys-
tem (MNS), comprising the inferior frontal gyrus and
inferior parietal cortex, has led to a surge of interest in
the neural correlates of social cognition, including
empathy. The broad notion that empathy involves
“putting oneself in another’s shoes,” by simulating
what others do, think, or feel, has been linked to the
properties of mirror neurons (Iacoboni & Mazziotta,
2007). Specifically, it has been proposed that the
“internal simulation” or “mirror matching” of actions
and emotions by the MNS occurs automatically and
unconsciously, and underlies our ability to empathize
(Carr, Iacoboni, Dubeau, Mazziotta, & Lenzi, 2003;
Gallese, 2003; Leslie, Johnson-Frey, & Grafton,
The adoption of mirror neurons as the neural
underpinning of empathy, or at least an integral part
of the neural network underlying this ability, is partly
based on a presumed link between imitation and
empathy. Surprisingly, there is limited empirical evid-
ence for this link. In this short review, we critically
evaluate studies that have examined the relationship
between imitation and empathy and their neural corre-
lates. We have chosen to focus our review on studies
published since 2008 to ensure that it is topical and
up-to-date. The role of the MNS in motor imitation
and action understanding has recently been questioned
(e.g., Hicock, 2008; Molenbergh, Cunnington, &
Mattingley, 2009), and evidence for its specific
involvement in empathic functioning is even less
Correspondence should be addressed to: Sarah J. Wilson, Psychological Sciences, University of Melbourne, Parkville, 3010, Australia.
This work was supported by funding from the Australian National Health and Medical Research Council (Project grant 566759) and a
Medical Research Grant from the University of Melbourne.
Downloaded by [ ] at 19:21 15 January 2012
328 BAIRD ET AL.
compelling (Decety, 2010). It is enticing to assert that
the MNS underpins empathy, and studies in several
different areas support this contention. These include
lesion studies of imitation and empathy, functional
neuroimaging showing a positive correlation between
empathy self-report and MNS activation, and obser-
vations of impaired imitation and empathy in autism
spectrum disorders (ASD). While valuable, these
studies do not as yet provide us with a comprehensive
framework for understanding the role of the MNS in
empathy, and researchers have recently challenged
the extent to which the MNS may be involved
As this review will show, research to date suggests
that there are various forms of imitation and empathy
that are likely to be underpinned by partially inde-
pendent neural substrates. These forms include emo-
tional and non-emotional, automatic, and voluntary
imitation, as well as motor, cognitive, and emotional
empathy (see summary in Table 1). Recent neuroim-
aging and lesion studies demonstrate that distinct neu-
ral regions underlie these forms, and each form recruit
the MNS to a variable degree (Bien, Roebroeck,
Goebel, & Sack, 2009; Shamay-Tsoory, Aharon-
Peretz, & Perry, 2008). Conceivably, the development
of a comprehensive account of empathy and its neural
underpinnings would need to address the relevance of
the relationships between these various forms and the
role of the MNS in each.
WHAT IS EMPATHY?
Theodor Lipps is regarded as the father of the concept
of einfühlung or empathy (translated from the German
by Tichener in 1909). Although Robert Vischer first
used the term in 1873 to describe the projection of
human feeling onto the natural world, Lipps (1903)
expanded this theory to explain how we understand
the mental states of others. He considered it to be an
unconscious process based on a “natural instinct” and
“inner imitation” (Montag, Gallinat, & Heinz, 2008).
Since then, empathy has been defined in many ways,
and definitional debates have predominated, including
whether empathy is a cognitive or an emotional
process (Preston & de Waal, 2002). For example,
some authors adopt a broad definition of empathy that
encompasses emotional contagion or sympathy,
while others prefer a narrower definition that distin-
guishes empathy from these related phenomena
(de Vignemont & Singer, 2006).
Different forms of empathy have been proposed.
Blair (2005) distinguished cognitive, motor, and emo-
tional empathy, and considered each to have a distinct
neural correlate. Others acknowledge various forms,
such as cognitive and emotional empathy, but argue
that in most instances these components co-occur and
cannot be disentangled (Baron-Cohen & Wheel-
wright, 2004). In addressing this issue, Preston and de
Waal (2002, p. 3) suggested that “the different views
of empathy can be cohered into a unified whole if a
broad view of the perception action model is taken.”
They conceived empathy as part of a group of related
processes, including imitation, which depend on a
“perception action mechanism.” These mechanisms
are thought to be underpinned by the same neural net-
work that mediates the perception and production of a
given behavior. This assumption is evident in recent
functional neuroimaging studies that have used imita-
tion paradigms as a proxy for an “empathy” condition
(see below). Thus, while differences in defining
empathy between studies make it difficult to compare
A schematic overview of the different forms of imitation and empathy and their main neural correlates identified by
research to date
Function Main neural correlates Type of research studies
Emotional Automatic Unknown No studies to date
Voluntary Right inferior frontal gyrus, “limbic” systemafMRI
Non-emotional Automatic Frontal regions: left frontal operculumbLesion, fMRI, TMS
Voluntary Frontal regions: premotor cortex, inferiorfrontal gyrus
Parietal regions: superior and inferiorparietal lobules
Lesion, fMRI, TMS
Motor “Sensorimotor” MNS TMS
Emotional Inferior frontal gyrus Lesion, fMRI
Cognitive Ventromedial prefrontal cortex Lesion, fMRI
fMRI: functional magnetic resonance imaging; TMS: transcranial magnetic stimulation; MNS: mirror neuron system.
aSee Iacoboni (2009). bSee Bien et al. (2009).
Downloaded by [ ] at 19:21 15 January 2012
MIRROR NEURON SYSTEM IN EMPATHY 329
findings, they nonetheless allow examination of MNS
involvement in various forms of empathy and imita-
tion. In this review, we will define the type of empa-
thy (corresponding with the measure used) that each
study investigated, where possible. When it is not
specified, the reader can assume that we are referring
to emotional empathy.
Measures of individual differences
Numerous measures of empathy have been previously
reviewed by Baron-Cohen and Wheelwright (2004).
Here, we briefly review only two self-report question-
naires that are relevant to the discussion of this review.
The Interpersonal Reactivity Index (IRI) (Davis,
1983) is the most commonly used questionnaire in
empathy research. This measure comprises four scales
of seven items each: (1) perspective taking (PT),
assessing cognitive empathy or the ability to under-
stand another’s point of view; (2) fantasy scale (FS),
measuring the ability to imagine oneself in the place of
fictional characters in books or movies; (3) empathic
concern (EC), assessing emotional empathy or
expressed concern for others; and (4) personal distress
(PD), measuring the emotional response induced by
observing strong emotions in others. This scale has
good internal consistency, with alpha coefficients
ranging from .68 to .79 (Davis, 1983). Although it has
been criticized as measuring processes broader than
empathy, such as self-control and imagination, it is
generally considered the most comprehensive measure
available (Baron-Cohen & Wheelwright, 2004).
The Questionnaire Measure of Emotional Empathy
(QMEE) (Mehrabian & Epstein, 1972) contains seven
subscales assessing the tendency to react to another’s
experience. Its split-half reliability is high (.84), sug-
gesting that it measures a single construct. The
authors propose this is emotional arousal to the envir-
onment in general, rather than the emotions of others
specifically. Thus, while the scale has been used to
assess affective empathy, it may be measuring a
different but related construct (Baron-Cohen &
THE RELATIONSHIP BETWEEN
IMITATION AND EMPATHY
Despite the long-held association between imitation and
empathy, there are few empirical studies of individual
differences in this relationship. Only two studies have
examined both abilities in healthy individuals. Chartrand
and Bargh (1999) investigated unconscious motor
mimicry, or what they termed the “chameleon effect,”
in participants during social interactions. They
described the mechanism underlying this effect as a
“perception-behavior link,” or the tendency to auto-
matically engage in behavior that one has perceived in
another. Chartrand and Bargh considered this a pas-
sive, cognitive process that is not associated with an
affective or emotional state. Correspondingly, they
predicted that individual differences in emotional
empathy would not modulate the chameleon effect. In
support of their prediction, they found a higher fre-
quency of motor imitation (foot shaking, face rubbing,
smiling) in individuals with higher cognitive empathy
scores (PT scale of the IRI) (Davis, 1983), but no
association between motor mimicry and emotional
empathy scores (EC scale of the IRI).
Sonnby-Borgström (2002) subsequently reported
findings that appear inconsistent with the work of
Chartrand and Bargh (1999). She used electromyogra-
phy (EMG) to measure the motor mimicry of emo-
tional facial expressions, such as happy and angry.
Individuals with high emotional empathy (as meas-
ured by the QMEE) showed a higher degree of facial
expression mimicry than those with low emotional
empathy. This led Sonnby-Borgström (2002) to con-
clude that “automatic mimicry is an early automatic
element of emotional empathy” (p. 439).
The findings of these studies suggest that the rela-
tionship between imitation and empathy may vary
depending on the particular form of each of these abil-
ities. Specifically, facial imitation was associated
with emotional empathy (Sonnby-Borgström, 2002),
while motor imitation was related to cognitive but not
emotional empathy (Chartrand & Bargh, 1999). It is
noteworthy, however, that Chartrand and Bargh’s def-
inition of motor imitation included smiling, a form of
facial imitation. The inconsistent results in relation to
facial imitation may be due to the use of different
stimuli (i.e., static photographs versus actors), or the
different measures of emotional empathy used in each
study. Further research is therefore required to clarify
the relationship between these different forms of
imitation and empathy, using a range of stimuli.
Neural correlates of imitation
Neuroimaging studies of empathy
Numerous neuroimaging studies have investigated
empathy with a variety of stimuli and paradigms, the
majority using observation of pain in others as a
Downloaded by [ ] at 19:21 15 January 2012
330 BAIRD ET AL.
model (Jackson, Rainville, & Decety, 2006). In gen-
eral, these studies have shown shared neural activa-
tion for observing and experiencing pain that includes
the bilateral anterior insula, dorsal anterior cingulate,
and sensorimotor cortices. Although these findings
appear to support the notion that empathy for pain
involves an automatically activated “sensorimotor
resonance” between self and other, Decety and col-
leagues (Decety, 2010; Yamada & Decety, 2009)
have recently argued that the involvement of these
brain regions may simply reflect a general aversive
response and activation of the threat detection and
defensive preparation system. Furthermore, studies of
disgust (Wicker et al., 2003) and taste (Jabbi, Swart, &
Keysers, 2007) demonstrate the integral role of the
insula in empathy, and thus highlight the importance
of regions outside the MNS in empathy for pain and
other specific emotions.
MNS involvement in empathy has been suggested
by four functional neuroimaging studies reporting a
positive correlation between MNS activation and
empathy scores derived from the IRI (Gazzola, Aziz-
Zadeh, & Keysers, 2006; Jabbi et al., 2007; Kaplan &
Iacoboni, 2006; Pfeifer, Iacoboni, Mazziotta, &
Dapretto, 2007). These studies are summarized in
Table 2 and show differing associations between spe-
cific IRI scales (and their corresponding forms of
empathy) and localized regions of MNS activation.
Three of the four studies found that MNS activation
correlated with emotional empathy (EC or PD scales)
(Jabbi et al., 2007; Kaplan & Iacoboni, 2006; Pfeifer
et al., 2007). These studies varied according to pas-
sive observation versus active imitation of the in-
scanner task, and included both auditory and visual
paradigms. The direction of the correlation was nega-
tive in one study (Kaplan & Iacoboni, 2006) and pos-
itive in two (Jabbi et al., 2007; Pfeifer et al., 2007). In
contrast, Gazzola et al. (2006) found a positive corre-
lation between cognitive empathy (PT scale) and the
MNS, specifically left parietal lobule activation.
Taken together, these studies are generally consid-
ered to support MNS involvement in emotional empa-
thy, most commonly seen as increased activation of
the inferior frontal gyrus either unilaterally or bilater-
ally. It is noteworthy, however, that all the studies
used out-of-scanner correlations, the significant pit-
falls of which have been outlined by Yarkoni (2009),
as well as by Vul, Harris, Winkielman, and Pashler
(2009). Methodological differences between these
studies also question the strength of the conclusions.
Although all the studies used the same out-of-scanner
measure of empathy, the in-scanner paradigms, type
of stimuli, and age of the participants varied widely
Moreover, prefrontal brain regions, such as the infe-
rior frontal gyrus, may be activated for reasons other
than an individual’s empathic abilities. As noted by
Cabeza and Nyberg (2000), this brain region is consist-
ently activated during a wide range of tasks, and its
engagement may reflect attention or linguistic proc-
esses rather than empathic functioning per se. Mirror
neurons account for only a minority of the cell popula-
tion in this area in the monkey, yet, despite this, Decety
(2010) has noted a bias in human neuroimaging
research to attribute a hemodynamic response in the
inferior frontal gyrus to MNS activity. Nevertheless, a
recent lesion study (Shamay-Tsoory et al., 2008) sup-
ports the preliminary neuroimaging findings of Table 2.
This study demonstrated dissociable neuroanatomical
substrates for cognitive and emotional empathy, with
emotional empathy deficits occurring after damage to
the inferior frontal gyrus, while impaired cognitive
empathy was manifested after ventromedial prefrontal
cortical damage (see section below). This finding again
highlights the need for a more comprehensive frame-
work that identifies the roles of specific brain regions in
the various forms of empathy.
Other functional neuroimaging studies have used
imitation paradigms as a proxy for empathy (e.g., Carr
et al., 2003); however, this approach is problematic.
Currently, there is limited empirical evidence of an
association between imitation and empathy. Given
that the MNS has been found to play a role in imita-
tion, these studies inevitably conclude that the MNS is
involved in empathy. As noted by Decety and Michal-
ska (2010, p. 896), “there is a problem with equating
empathy with motor resonance because the latter does
not convey insight into another’s internal state and
does not account for any other-oriented motivational
state that characterises sympathy.”
A distinction has also been typically made between
voluntary (conscious) and automatic (unconscious)
imitation, with the latter assumed to underlie
empathic functioning (Leslie et al., 2003). Voluntary
imitation typically requires the experimenter to ask
the participant to copy an action or expression. Per-
formance on such tasks may be influenced by nonspe-
cific factors such as executive function (Bird,
Leighton, Press, & Heyes, 2007). Automatic imitation
requires a participant to observe an action or expres-
sion, either passively or accompanied by a simple
movement, while the experimenter measures involun-
tary movements or the performance speed of the
simple movement. In this way, automatic imitation is
considered a purer measure of imitative ability.
Despite this, to date there have been no functional
neuroimaging studies investigating empathy that use
an automatic imitation task. This is a significant gap
Downloaded by [ ] at 19:21 15 January 2012
Summary of neuroimaging studies showing a positive correlation between self-reported empathy and MNS activation
Author (year) Participants (mean age) Imaging task and stimuli Empathy measure Main findings
Kaplan et al. (2006) 26 adults (26 years) Observation of film clips of drinking cups
in different contexts (cleaning, drinking,
and no context) and with different hand
grips (precision or whole hand)
Interpersonal Reactivity Index
(IRI)* (Davis, 1983)
Positive correlation between right inferior frontal gyrus
activation and empathic concern scale (average across
all film clips), and fantasy scale scores (during
incongruent and context alone clips)
Negative correlation with personal distress scale scores
(during congruent actions and action in context clips)
Gazzola et al. (2006) 16 adults (31 years) Listening to or performing mouth or hand
action sounds (e.g., kissing, ripping paper)
IRI Positive correlation between left premotor cortex and inferior
parietal lobule activation (during hand and environmental
sounds) and perspective-taking scale scores
(Note that participants were categorized into high and low
Jabbi et al. (2007) 18 adults (24 years) Observation of film clips of actors drinking
liquids and displaying pleased, disgusted,
and neutral facial expressions, and actual
ingestion of pleasant, disgusting, and
neutral tasting liquids
IRI Positive correlation between bilateral anterior insula and frontal
operculum activation and total IRI score (particularly evident
for the personal distress and fantasy scale scores)
No correlation between IRI scores and brain activation during
Pfeifer et al. (2008) 16 children (10 years) Imitation and observation of five facial
expressions (happy, fearful, sad, angry,
IRI and Interpersonal
Competence Scale (parental
Positive correlation between inferior frontal gyrus activation (in
addition to right insula, left amygdala, and left fusiform gyrus)
and IRI scale scores (except perspective-taking scale) during
observation and imitation of facial expressions
*See section on measures of empathy for a description of this questionnaire.
Downloaded by [ ] at 19:21 15 January 2012
332 BAIRD ET AL.
in our current knowledge. Future neuroimaging and
lesion studies will need to distinguish between auto-
matic and voluntary imitation involving different
body parts (i.e., face, hand, body) and determine how
they relate to the various types of empathy and their
While recent neuroimaging research has focused
on the neural networks associated with imitation and
empathy, only lesion or transcranial magnetic stimu-
lation (TMS) studies can identify brain regions that
are necessary for these functions. The majority of
neuropsychological studies examining the neural
basis of imitation and empathy have involved patients
with frontal lobe lesions, who typically show
enhanced imitation or impaired empathy (e.g., Brass,
Derrfuss, Mathes-von Cramon, & von Cramon, 2003;
Eslinger, 1998; Luria, 1980; Shamay-Tsoory et al.,
2008). Damage to the left parietal region has also
been shown to alter imitation performance (Halsband
et al., 2001), while there appear to be no studies
reporting impaired emotional processing in these
patients. In general, lesion studies suggest a “core cir-
cuitry of imitation” underpinned by the MNS; how-
ever, human lesion research has yet to identify more
specific areas within frontal and parietal regions.
Notably, a recent meta-analysis of 20 functional MRI
studies that examined the role of frontal and parietal
brain regions in imitation suggested that areas extend-
ing beyond the classic MNS are also crucial for imita-
tion (Molenbergh et al., 2009).
Two recent TMS studies using self-report question-
naires have demonstrated an association between “sen-
sory” or “motor” empathy for pain and MNS function
(Avenanti, Bueti, Galati, & Aglioti, 2005; Fecteau,
Pascual-Leone, & Theoret, 2008). Both studies meas-
ured motor cortex excitability as a marker of “sensorim-
otor” MNS function. Although the motor cortex is not
typically considered part of the human MNS (Iacoboni
& Mazziotta, 2007), mirror neurons have been found in
the monkey primary motor cortex (Tkach, Reimer, &
Hatsopoulos, 2007). The findings of these two TMS
studies provide support for the motor system being
involved specifically in awareness of pain.
ASD: ARE IMPAIRED IMITATION AND
EMPATHY A REFLECTION OF MNS
From observations of impaired imitation and
empathic abilities in individuals with ASD, it has
been hypothesized that the MNS plays a critical role
in these deficits (Williams, Whiten, Suddendorf, &
Perrett, 2001). Recent studies, however, have chal-
lenged the presence of either a global imitation or an
empathic deficit in ASD.
Imitation in ASD
The first observation of an imitative deficit in autism
was made over 50 years ago (Ritvo & Provence,
1953). Since then, there have been numerous studies
of imitation in the ASD population, the majority
focusing on motor imitation (meaningful/meaningless
gestures or actions). In general, these studies show
impaired imitative abilities in ASD compared with
controls (Williams, Whiten, & Singh, 2004).
More recently, three studies have demonstrated
atypical activation of MNS regions during perform-
ance of various motor imitation tasks in ASD groups
compared with controls (Dapretto et al., 2006; Nishi-
tani, Avikainen, & Hari, 2004; Williams et al., 2006).
These findings have been cited as support for MNS
dysfunction in ASD. The issue here, however, is that
performance of the imitation task by the ASD group
was intact (Dapretto et al., 2006; Nishitani et al.,
2004; Williams et al., 2006). This suggests that atypi-
cal MNS activation does not necessarily reflect MNS
dysfunction per se, as imitation was not impaired.
Rather, individuals with ASD may simply recruit
alternative brain regions or use the MNS differently to
perform such tasks. All three studies measured volun-
tary imitation, which may be dissociated from auto-
matic imitative abilities in ASD (McIntosh,
Reichmann-Decker, Winkielman, & Wilbarger, 2006;
Tardif, Laine, Rodriguez, & Gepner, 2007). Only a
few studies have examined automatic imitation in the
ASD population, with inconsistent results. Bird et al.
(2007) found intact automatic imitation of robotic and
human hand actions in adults with ASD. Moreover,
Tardif et al. (2007) found enhanced automatic imita-
tion of facial and vocal stimuli in children with ASD
compared with controls.
In general, the differing imitation results in ASD
probably reflect methodological variations, such as
stimulus type (motor or facial/emotional), the age of
participants, and the method of measuring automatic
imitation (behavioral observations or physiological
measures such as EMG). Equally plausible, however,
is the suggestion that imitation may not be a unitary
construct, but may depend on more than one neural
network. Hamilton, Brindley, and Frith (2007), using
“mirror neuron” tasks previously employed in
neuroimaging studies, reported intact hand action
Downloaded by [ ] at 19:21 15 January 2012
MIRROR NEURON SYSTEM IN EMPATHY 333
imitation in children with ASD. They suggested that
different imitation behaviors may be supported by dif-
ferent neural systems. They also reported impaired
“theory of mind” (cognitive empathy) in this sample.
Accordingly, various types of empathy may be associ-
ated with different forms of imitation, and the integ-
rity of these different forms and their relationships to
cognitive and emotional empathy requires systematic
investigation in both healthy and ASD populations.
Empathy in ASD
Impaired empathy is considered a hallmark feature of
ASD (Baron-Cohen & Wheelwright, 2004; Blair,
2008; Gillberg, 1992). Despite this, there is a paucity
of research on the nature of empathic functioning in
ASD, particularly in relation to specific forms of
empathy. Extant research has primarily focused on
one form of empathy, namely cognitive empathy, the
majority of studies showing theory of mind deficits.
This focus has led to the conclusion that empathy is
impaired in ASD. Recent studies, however, have
begun to adopt a broader framework, challenging the
notion that empathy is a unitary concept by examining
different forms of empathy in this population.
Three studies have assessed emotional empathy in
the ASD population to date. In a case study of two
individuals with Asperger syndrome (AS), Shamay-
Tsory, Tomer, Yaniv, and Aharon-Peretz (2002),
using the IRI and QMEE, found that both cognitive
and emotional empathy were impaired. Two group
studies, however, refute this finding. Rogers, Dzi-
obek, Hassenstab, Wolf, and Convit (2007) used the
IRI to assess empathy in a sample of individuals with
AS and found that while cognitive empathy was
impaired, scores on the emotional empathy scale were
similar to controls. This finding was replicated by
Dziobek et al. (2008). These findings question
whether all forms of empathy are impaired in ASD
and raise the possibility that cognitive and emotional
forms of empathy may be dissociated in this popula-
tion. This notion is supported by the findings of the
lesion study by Shamay-Tsoory et al. (2008) described
above. It poses a challenge for the MNS dysfunction
hypothesis of ASD, which assumes that empathy is a
unitary phenomenon underpinned by the MNS.
In this paper, we have critically examined the current
evidence for MNS involvement in empathy. The
assumption of an intimate relationship between imitation
and empathy has contributed to the notion of the MNS
as the neural correlate of empathy. There is, however,
limited empirical evidence for such an association.
Further research is required to clarify the exact nature
of this relationship in both healthy and neurologically
impaired populations. In our review of functional neu-
roimaging studies showing positive correlations
between MNS activation and empathy scale scores,
we found that the results are mixed, despite use of the
same behavioral measure of empathic functioning
(the IRI). While MNS dysfunction has been demon-
strated in ASD, undoubtedly complex links exist
between the nature of the imitative or empathic deficit
and the network involved. To summarize, we propose
that a more comprehensive framework that promotes
systematic investigation of the various forms of
empathy is needed to guide future research. Develop-
ment of this framework offers the opportunity to
extend our understanding of the neural bases of this
complex and fundamental human ability.
Manuscript received 17 February 2010
Manuscript accepted 25 November 2010
First published online 10 January 2011
Avenanti, A., Bueti, D., Galati, G., & Aglioti, S. M.
(2005). Transcranial magnetic stimulation highlights
the sensorimotor side of empathy for pain. Nature Neu-
roscience, 8, 955–960.
Baron-Cohen, S., & Wheelwright, S. (2004). The empathy
quotient: An investigation of adults with Asperger syn-
drome or high functioning autism, and normal sex differ-
ences. Journal of Autism and Developmental Disorders,
Bien, N., Roebroeck, A., Goebel, R., & Sack, A. T. (2009).
The brain’s intention to imitate: The neurobiology of
intentional versus automatic imitation. Cerebral Cortex,
Bird, G., Leighton, J., Press, C., & Heyes, C. (2007). Intact
automatic imitation of human and robot actions in
autism spectrum disorders. Proceedings of the Royal
Society. Series B: Biological Sciences, 274, 3027–3031.
Blair, R. (2008). Fine cuts of empathy and the amygdala:
Dissociable deficits in psychopathy and autism.
Quarterly Journal of Experimental Psychology, 61,
Blair, R. J. (2005). Responding to the emotions of others:
Dissociating forms of empathy through the study of typi-
cal and psychiatric populations. Consciousness and Cog-
nition, 14, 698–718.
Brass, M., Derrfuss, J., Matthes-von Cramon, G., & von
Cramon, D. Y. (2003). Imitative response tendencies in
patients with frontal brain lesions. Neuropsychology, 17,
Cabeza, R., & Nyberg, L. (2000). Imaging cognition II: An
empirical review of 275 PET and fMRI studies. Journal
of Cognitive Neuroscience, 12, 1–47.
Downloaded by [ ] at 19:21 15 January 2012
334 BAIRD ET AL.
Carr, L., Iacoboni, M., Dubeau, M. C., Mazziotta, J. C.,
& Lenzi, G. L. (2003). Neural mechanisms of empa-
thy in humans: A relay from neural systems for imita-
tion to limbic areas. Proceedings of the National
Academy of Sciences of the United States of America,
Chartrand, T. L., & Bargh, J. A. (1999). The chameleon
effect: The perception-behavior link and social interac-
tion. Journal of Personality and Social Psychology, 76,
Dapretto, M., Davies, M. S., Pfeifer, J. H., Scott, A. A., Sig-
man, M., Bookheimer, S. Y., & Iacoboni, M. (2006).
Understanding emotions in others: Mirror neuron dys-
function in children with autism spectrum disorders.
Nature Neuroscience, 9, 28–30.
Davis, M. (1983). Measuring individual differences in
empathy: Evidence for a multidimensional approach.
Journal of Personality and Social Psychology, 44,
Decety, J. (2010). To what extent is the experience of empa-
thy mediated by shared neural circuits? Emotion Review,
Decety, J., & Michalska, K. J. (2010). Neurodevelopmental
changes in the circuits underlying empathy and sympathy
from childhood to adulthood. Developmental Science, 13,
de Vignemont, F., & Singer, T. (2006). The empathic brain:
How, when and why? Trends in Cognitive Science, 10,
Dziobek, I., Rogers, K., Fleck, S., Bahnemann, M., Heek-
eren, H. R., Wolf, O. T., et al. (2008). Dissociation of
cognitive and emotional empathy in adults with
Asperger syndrome using the Multifaceted Empathy
Test (MET). Journal of Autism and Developmental
Disorders, 38, 464–473.
Eslinger, P. J. (1998). Neurological and neuropsychological
bases of empathy. European Neurology, 39, 193–199.
Fecteau, S., Pascual-Leone, A., & Theoret, H. (2008). Psy-
chopathy and the mirror neuron system: Preliminary
findings from a non-psychiatric sample. Psychiatry
Research, 160, 137–144.
Gallese, V. (2003). The roots of empathy: The shared mani-
fold hypothesis and the neural basis of intersubjectivity.
Psychopathology, 36, 171–180.
Gazzola, V., Aziz-Zadeh, L., & Keysers, C. (2006). Empa-
thy and the somatotopic auditory mirror system in
humans. Current Biology, 16, 1824–1829.
Gillberg, C. (1992). The Emanuel Miller Memorial Lecture
1991. Autism and autistic like conditions: Subclasses
among disorders of empathy. Journal of Child Psychol-
ogy and Psychiatry, 33, 813–842.
Halsband, U., Schmitt, J., Weyers, M., Binofski, F., Grut-
zner, G., & Freund, H.-J. (2001). Recognition and imita-
tion of pantomimed motor acts after unilateral parietal
and premotor lesions: A perspective on apraxia. Neu-
ropsychologia, 39, 200–216.
Hamilton, A. F., Brindley, R. M., & Frith, U. (2007). Imita-
tion and action understanding in autistic spectrum disor-
ders: How valid is the hypothesis of a deficit in the mirror
neuron system? Neuropsychologia, 45, 1859–1868.
Hicock, G. (2008). Eight problems for the mirror neuron
theory of action understanding in monkeys and humans.
Journal of Cognitive Neuroscience, 21, 1229–1243.
Iacoboni, M. (2009). Imitation, empathy, and mirror neu-
rons. Annual Review of Psychology, 60, 653–670.
Iacoboni, M., & Mazziotta, J. C. (2007). Mirror neuron sys-
tem: Basic findings and clinical applications. Annals of
Neurology, 62, 213–218.
Jabbi, M., Swart, M., & Keysers, C. (2007). Empathy for
positive and negative emotions in the gustatory cortex.
NeuroImage, 34, 1744–1753.
Jackson, P., Rainville, P., & Decety, J. (2006). To what
extent do we share the pain of others? Insights from the
neural bases of pain empathy. Pain, 125, 5–9.
Kaplan, J. T., & Iacoboni, M. (2006). Getting a grip on
other minds: Mirror neurons, intention understanding,
and cognitive empathy. Society for Neuroscience, 1,
Leslie, K., Johnson-Frey, S., & Grafton, S. (2003). Func-
tional imaging of face and hand imitation: Towards a
theory of empathy. NeuroImage, 21, 601–607.
Lipps, T. (1903). Einfühlung, innere Nachahmung und
Organempfindung. für gesamte Psychologie, 1, 465–
519. Trans. as: Empathy, inner imitation and sense-feel-
ings. In M. Rader (Ed.), A modern book of esthetics
(1979; 5th ed., pp. 374–382). New York, NY: Holt,
Rinehart and Winston.
Luria, A. R. (1980). Higher cortical functions in man. New
York, NY: Basic Books.
McIntosh, D. N., Reichmann-Decker, A., Winkielman, P.,
& Wilbarger, J. L. (2006). When the social mirror
breaks: Deficits in automatic, but not voluntary, mimicry
of emotional facial expressions in autism. Developmen-
tal Science, 9, 295–302.
Mehrabian, A., & Epstein, N. (1972). A measure of emo-
tional empathy. Journal of Personality, 40, 525–543.
Molenbergh, P., Cunnington, R., & Mattingley, J. B. (2009).
Is the mirror system involved in imitation? A short
review and meta-analysis. Neuroscience and Biobehav-
ioral Review, 33, 975–980.
Montag, C., Gallinat, J., & Heinz, A. (2008). Theodor Lipps
and the concept of empathy: 1851–1914. American
Journal of Psychiatry, 165, 1261.
Nishitani, N., Avikainen, S., & Hari, R. (2004). Abnormal
imitation-related cortical activation sequences in
Asperger’s syndrome. Annals of Neurology, 55, 558–562.
Pfeifer, J. H., Iacoboni, M., Mazziotta, J. C., & Dapretto, M.
(2007). Mirroring others’ emotions relates to empathy
and interpersonal competence in children. NeuroImage,
Preston, S. D., & de Waal, F. B. (2002). Empathy: Its ultimate
and proximate bases. Behavioral and Brain Sciences,
Ritvo, S., & Provence, S. (1953). Form perception and imi-
tation in some autistic children: Diagnostic findings and
their contextual interpretation. The Psychoanalytic Study
of the Child, 8, 155–161.
Rogers, K., Dziobek, I., Hassenstab, J., Wolf, O. T., & Con-
vit, A. (2007). Who cares? Revisiting empathy in
Asperger syndrome. Journal of Autism and Develop-
mental Disorders, 37, 709–715.
Shamay-Tsoory, S. G., Aharon-Peretz, J., & Perry, D.
(2008). Two systems for empathy: A double dissociation
between emotional and cognitive empathy in inferior
frontal gyrus versus ventromedial prefrontal lesions.
Brain, 132, 617–627.
Downloaded by [ ] at 19:21 15 January 2012
MIRROR NEURON SYSTEM IN EMPATHY 335
Shamay-Tsoory, S. G., Tomer, R., Yaniv, S., & Aharon-
Peretz, J. (2002). Empathy deficits in Asperger syn-
drome: A cognitive profile. Neurocase: Case Studies in
Neuropsychology, Neuropsychiatry, and Behavioural
Neurology, 8, 245–252.
Sonnby-Borgström, M. (2002). Automatic mimicry reac-
tions as related to differences in emotional empathy.
Scandinavian Journal of Psychology, 43, 433–443.
Tardif, C., Laine, F., Rodriguez, M., & Gepner, B. (2007).
Slowing down presentation of facial movements and vocal
sounds enhances facial expression recognition and induces
facial-vocal imitation in children with autism. Journal of
Autism and Developmental Disorders, 37, 1469–1484.
Tkach, D., Reimer, J., & Hatsopoulos, N. (2007). Congruent
activity during action and action observation in motor
cortex. Journal of Neuroscience, 27, 13241–13250.
Vul, E., Harris, C., Winkielman, P., & Pashler, H. (2009).
Puzzlingly high correlations in fMRI studies of emotion,
personality and social cognition. Perspectives on Psy-
chological Science, 4, 274–290.
Wicker, B., Keysers, C., Piailly, J., Royet, J.-P., Gallese, V.,
& Rizzolatti, G. (2003). Both of us disgusted in my
insula: The common neural basis of seeing and feeling
disgust. Neuron, 40, 655–664.
Williams, J. H., Waiter, G. D., Gilchrist, A., Perrett, D.
I., Murray, A. D., & Whiten, A. (2006). Neural mech-
anisms of imitation and ‘mirror neuron’ functioning in
autistic spectrum disorder. Neuropsychologia, 44,
Williams, J. H., Whiten, A., & Singh, T. (2004). A system-
atic review of action imitation in autistic spectrum disor-
der. Journal of Autism and Developmental Disorders,
Williams, J. H., Whiten, A., Suddendorf, T., & Perrett, D. I.
(2001). Imitation, mirror neurons and autism. Neuro-
science and Biobehavioral Reviews, 25, 287–295.
Yamada, M., & Decety, J. (2009). Unconscious processing
and empathy: An investigation of subliminal priming on
the detection of painful facial expressions. Pain, 143,
Yarkoni, T. (2009). Big correlations in little studies. Inflated
fMRI correlations reflect low statistical power-commen-
tary on Vul et al. (2009). Perspectives on Psychological
Science, 4, 294–298.
Downloaded by [ ] at 19:21 15 January 2012