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The contribution of transcranial magnetic stimulation (TMS) to our understanding of the

processes underlying empathy.

Transcranial magnetic stimulation (TMS) is a noninvasive technique whereby nerve cells in the
brain are stimulated using electromagnetic fields delivered through a coil held against the head
(Mayo Clinic, 2015). It is primarily used as a method of treating depression, however it is
increasingly being used as a research method in cognitive neuropsychology. Over the last decade,
researchers have been using TMS to study empathy; the process of understanding, and sometimes
experiencing, what others are thinking or feeling (Pijnenborg, Spikman, Jeronimus, Aleman, 2012).
Hetu, Taschereau-Dumouchel, and Jackson (2012) identify three distinct components of empathy;
resonance; a bottom-up process whereby brain activity is influenced by observing the affective state
of someone else, mentalizing; a deliberate top-down process allowing us to understand the mental
states and intentions of others, and self-other discrimination; the ability to attribute the source of an
affective state to oneself or others. This essay will review some of the studies which have used TMS
to study these processes underlying empathy. I will discuss the merits of using TMS to study
empathy, in contrast to other methods, and consider possible directions for future research
throughout.
Resonance has primarily been studied through the phenomena of sensorimotor contagion; the
reduction of corticospinal excitability which occurs when observing somebody else experiencing
pain (Farina, Tinazzi, Le Pera, & Valeriani, 2003). A study by Avenanti, Bueti, Galati and Aglioti
(2005) was one of the first to use TMS to explore this sensorimotor aspect of empathy. Peyron,
Laurent, and Garcia-Larrea (2000) proposed that pain is represented in a corticosubcortical network
called the Pain Matrix which comprises sensorimotor (representing location and intensity) and
affective (representing unpleasentness) nodes. Previous research (Singer, Seymour, ODoherty,
Kaube, Dolan, & Frith, 2004) indicates that only the affective component of the pain network is
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involved in empathy (i.e. the anterior cingulate cortex and the anterior insula). However, using
TMS, Avenanti et al. were able to show that perceiving pain in others was associated with a
significant drop in corticospinal excitability in the sensorimotor system.
Applying a strong magnetic pulse to specific areas of the motor cortex produces motor evoked
potentials (MEPs) in the associated muscles. The strength of the TMS pulse needed to evoke MEPs
is an inidcator of corticospinal excitability. As with real pain, seeing others experience pain prompts
a specific corticospinal inhibition, suggesting the activation of pain representations in the observers
sensorimotor system. TMS was used to measure changes in corticospinal motor representations for
the hand muscles of participants who were asked to watch the hands or feet of a human model or
noncorporeal objects being penetrated by a needle. Participants in the experimental condition of this
study watched a video of a needle being pushed into a hand, whilst those in the control conditions
watched a cotton bud pressing the hand or a needle being pushed into a tomato. The results show a
significant drop in the magnitude of MEPs specific to the muscle that participants saw being
pricked (i.e., participants seeing a particular hand muscle being pricked with a needle experienced a
reduction of motor excitability in their own corresponding muscle). No change in corticospinal
excitability occurred in the control conditions, indicating that the reduced excitability was
associated with observing the pain of another person.
MEPs were recorded from the first dorsal interosseous (FDI, in the index finger) and the abductor
digiti minimi (ADM, in the pinky finger) muscles of the right hands of participants. Watching a
needle penetrating a models foot did not affect the corticospinal excitability of these muscles.
MEPs measured at the FDI muscle of the participants were prompted by seeing the needle enter the
FDI muscle of the models hand but not the ADI muscle. MEPs measured from participants ADM
muscle demonstrated the opposite pattern. This inhibition of excitability correlated significantly
with participants subjective rating of the models pain. It also correlated with measures of sensory,
but not emotional empathy. These results indicate that embodying an understanding of other
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poeples pain into the motor system is important for pain-related social learning.
Singer and Frith (2005) provide possible explanations for how Avenanti et al. found a
sensorimotor empathic response when previous studies did not. This may be due to the different
materials used to evoke empathy. Participants in Avenanti et als study saw a strangers hands being
pricked with a needle likely emphasising the sensory aspect of the pain. Conversely, in Singer et als
2004 study participants were shown an arrow to indicate when their partner would experience a
painful stimulus likely emphasising the affective aspect of the pain.
However even functioning magnetic resonance imaging (fMRI) studies (Jackson, Meltzoff, &
Decety, 2005) that use materials highlighting the sensorimotor side of pain found different results.
This may be because TMS can pick up on slight changes in the sensorimotor system that are below
the significance threshold for fMRI, which does not always detect activation in the somatosensory
cortex (SI), even when participants receive painful stimuli. According to one meta-analysis (Peyron
& Laurent, 2000), only 50% of imaging studies examining pain reported activity in the SI. These
discrepancies have also been observed in action observation research. TMS studies have shown that
action observation can change corticospinal excitation and may be mapped directly to the specific
muscles used (Fadiga, Craighero, & Olivier, 2005). Conversly, fMRI studies of action observation
usually show activity in the inferior frontal gyrus and the inferior parietal lobule, moreso than the
primary sensorimotor cortex (Rizzolatti & Craighero, 2004).
However it is likely that there is more to empathy than the neural mapping of the sensory and
affective aspects of others pain. Individual differences in emotional and cognitive empathy are
likely to influence reaction to witnessing pain. Lamm, Batson, and Decety (2007) recognised that
empathic reactions to the pain of others can be more other-oriented (expressed through concern for
others) or self-oriented (experiencing distress when witnessing pain). Whilst these two components
may work alongside one another, they have different implications for how the observer responds.
Another important aspect of empathy is the ability to understand the perspectives of others. This is
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known as cognitive empathy and entails thinking about and imagining the feelings of others,
without necessarily having an affective reaction (Davis, 1996). Avenanti et al. (2009) conducted a
study to research how differences in these distinct components of empathy modulate somatomotor
responses. This study followed a procedure similar to Avenanti et al. (2005) with participants
watching a needle penetrating a models hand whilst having TMS-prompted MEPs recorded. They
also included self-report measures for the emotional and cognitive aspects of empathy. As with the
previous study, a reduction in corticospinal excitability was observed specific to the muscle
observed (FDI) in the index finger. The somatomotor response was higher for participants who
reported higher levels of cognitive empathy and lower for participants who reported higher levels of
distress. Each of these measures predicted somatomotor pain responses independently, indicating
that the processes underlying empathy are associated in different ways within the sensorimotor
system. The results indicate that an increased tendency to cognitively simulate the affective states of
others strengthens the somatomotor response to pain through a top-down process and operates
independently of the emotional mechanism. An increased tendency to experience distress at the pain
of others seemed to correlate with a facilitation of corticospinal activity, and may lessen or possibly
block mapping the pain of others in the somatomotor system. This is supported by another TMS
study showing that observation of emotional stimuli causes an increase in corticospinal excitability
(Hajcak et al., 2007). The authors suggest that a reduced empathic response caused by personal
distress may be linked with a reduction in mirror-matching the mental states of others. This study
takes account of the variability in empathic reactivity to the pain of others.
We might note an alternative explanation of the observed effect through the activation of the
motor mirror system. Corticospinal inhibition when witnessing pain may indicate a defensive motor
reaction similar to a withdrawal reflex (Farina et al. 2003). However, such motor reactions tend to
involve MEP suppression in all distal hand muscles (Urban et al. 2004) and given the precision of
the effect observed in this experiment (there was no inhibition in the ADM muscle), a mass reflex
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activation is unlikely.
Further research has used TMS to explore how mentalizing activities, in this case racial bias, can
influence pain resonance. Previous research in social psychology has suggested that racial
discrimination often results from a lack of empathy (Feagin, Vera, & Batur, 2001), however little
research had been conducted into differential sensorimotor reactivity to the suffering of someone of
a similar or different racial background until a study by Avenanti, Sirigu, and Aglioti (2010). TMS
was used to explore the sensorimotor empathic brain responses in participants who demonstrated
implicit but not explicit in-group preference. Participants watched a video showing the right hand of
a black or white model being penetrated by a needle or being touched by a cotton bud. As with the
previous studies, the specific location was the first dorsal interosseous (FDI) of the right hand.
MEPs were recorded from the right FDI (target) and ADM (control site) hand muscles of the
participants. It was found that participants' corticospinal motor system was inhibited when
observing the pain of in-group models and of models with violet skin (indicating ethnic neutrality).
While this inhibition was specific to the FDI muscle, no such inhibition was recorded in the ADM.
Corticospinal inhibition was significantly lower when viewing the pain of out-group models, and
this reduction was positively correlated with scores of implicit racial bias (measured using an
adapted version of the Implicit Association Test). These results indicate that while people may have
an automatic empathetic response to other peoples suffering, this reaction can be inhibited by racial
stereotypes and biases resulting in a lower sensorimotor response.
It is possible that the observed effect is merely due to visual unfamiliarity or increased perceived
dissimilarity between participants and the model, as opposed to racial bias. However, given that
corticospinal reactivity to the violet hand, which was rated as the least familiar skin colour among
participants, was significantly stronger than the response to the outgroup hand, this is unlikely to be
the case. These results provide a neural basis for the idea that racial biases can influence social
categorisation, leading to a devalued view of out-group strangers. It also demonstrates an
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interaction between two components of empathy, with the mentalizing effect of racial bias
influencing resonance.
Another form of mentalizing acivity which has been studied with TMS is mental state attribution:
the ability to infer the mental states of others. fMRI studies have shown that there is increased
activity in the right temporoparietal junction (RTPJ) when reading about someone's beliefs in a
moral situation (Young, Cushman, Hauser, & Saxe, 2007). The functional selectivity of the RTPJ
for moral beliefs increases significantly between the ages of six and eleven. Compared to adults,
young children, whose ability to reason about other peoples mental states is not fully developed,
place a higher value on the consequences of actions, rather than intentions, in order to make moral
judgments (Karniol, 1978). However, fMRI is unable to say whether activity in this area is a
necessary condition for mental state reasoning or even moral judgment itself. Young, Camprodon,
Hauser, Pascual-Leone, and Saxe (2010) hypothesised that the RTPJ is necessary for making moral
judgments and that disruption in this area would lead participants to rely less on the mental states or
beliefs of an actor and place more importance on outcomes. In this study, TMS was applied to the
RTPJ in the experimental condition, and the right parietal cortex in the control condition. TMS was
applied while participants had to make a moral judgement about a scenario where the agent either
intends to harm somebody or not and either succeeds in harming them or does not (4 conditions).
The results show a significant interaction between TMS site and agent belief-state but not between
TMS site and outcome. Following TMS to the RTPJ, participants relied more on the outcomes of
the action and less on the agents mental state to judge the rightness or wrongness of that action.
Participants in the experimental condition rated attempted harms (the intention but failure to harm)
as more or less desirable than participants in the control condition. This implies that disrupting the
activity in the RTPJ interferes with individuals capacity to assimilate mental states when making
moral judgments, particularly with regard to attempted harms, but does not disrupt the process of
moral judgment-making altogether.
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It is notable that TMS to the RTPJ did not affect judgments of intentional harms (i.e. a harmful
action was not considered anymore immoral when the protagonist intended to cause harm, than
when the harm was accidental). Based on this, it is likely that moral judgments reflect the weighting
of other factors, such as outcome, when information about the agent's mental state is unavailable.
An alternative hypothesis is that RTPJ disruption impairs the very process of moral judgment,
particularly when there are multiple factors to consider. According to this hypothesis, participants
perform less well when assimilating information about multiple morally relevant considerations
(e.g., someones past history, the methods used, and any constraints on them) following TMS to
RTPJ. The authors reject this hypothesis since it would not predict the effects found in this
experiment. If the ability to take account of multiple morally relevant considerations was hindered
by TMS to the RTPJ then we would expect judgments to have been otherwise slower in the
experimental condition. Further, the results showed a systematic bias, in accordance with the
predictions for each condition, not resulting from a mere slowing process. Therefore, the authors
conclude that TMS to the RTPJ disrupts input to moral judgment making, such as information about
mental states, but does not affect the process of moral judgment making itself. However, the
interference of multiple morally relevant features on moral judgment would benefit from further
research.
Another possible explanation is that TMS to the RTPJ interfered with other cognitive processes.
The lateral inferior parietal region, an area of the brain associated with attention shifting, is located
near the RTPJ (Mitchell, 2007). However, a study by Decety and Lamm (2007) found that these two
regions are separated by approximately 10mm, whilst TMS has a spatial resolution of 5 to 10 mm
(Kammer, 1999). The particular region of the RTPJ associated with mental state attribution was
located using image-guided TMS and a functional localizer. Furthermore, the results of this
experiment do not suggest there was any interference with attention or any other effect on task
performance; participants judgments were no slower or less reliable (i.e., more or less variable)
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during TMS to the RTPJ or control site in any of the conditions. These results support the initial
hypothesis that the effect of TMS on the RTPJ is due to the stimulus itself (i.e., belief information).
Through the use of action observation and pain perception models, these TMS studies provide
good evidence supporting the contention that the resonance process gives rise to the sesnsoriaffective components of empathy. However, if these resonance responses use similar neural
pathways to the ones used for our own sensor-affective responses then there are likely to be neural
mechanisms which facilitate the identification of the source of our internal state, (Hetu et al. 2012).
The process of determining whether an affective internal state arises from oneself or from
observing others is referred to as self-other discrimination (Hetu et al. 2012). Several studies have
been conducted whereby rTMS is applied during agency attribution or self-other discrimination
tasks to discover which areas of the brain are associated with this process. David et al. (2009)
identified the extrastriate body area (EBA) as a brain region instrumental with this component of
empathy, given that performance on a self-other discrimination task deteriorates following rTMS
over the EBA. Notably, the tasks in this study required participants to judge whether the movements
of visually presented stimuli were caused by their own actions or the actions of somebody else.
Whilst this study may inform us about the self-other distinction, its use of limb movements may
place disproportionate focus on the motor aspects of these stimuli whilst facial expressions and
characteristics are more likely to be better indicators of a persons affective state.
Uddin et al. (2006) therefore asked participants, following a single session of rTMS to the right
inferior parietal lobule (IPL), to judge whether a face was theirs or that of someone they knew well.
The face of each participants was merged with the face of the other person to varying ratios of self/
other (e.g. 40% self to 60% other). The results indicated that task performance was significantly
impaired following rTMS to the IPL but not the control site (left IPL). In particular, there was a
higher number of self responses even when the face was comprised mostly of other
characteristics. These results may be interpreted as a less liberal technique in response to features of
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the other or the overinclusion of stimuli with features that are associated with the self (i.e., a
lower threshold for self criteria). Based on these results, Uddin et al. proposed that selfrecognition is part of a larger system that represents the self and serves two roles. They propose that
it establishes representations between the self and others to facilitate communication, and also that
it distinguishes representations of self from others. They cite developmental studies showing a link
between the emergence of self-recognition and an increase in imitation abilities (Asendorpf &
Baudonniere, 1993). However such co-occurring developments ought to be researched further to see
if they are causally linked. One limitation of this study is the use of static faces as stimuli given the
typical association of mirror neutron activation and action observation. However research
demonstrating the activation of mirror neurons in response to static stimuli, whilst minimal, does
imply a similar process may be happening in this study. Given that this study did not include a
control condition involving discriminating between a non-self face and a familiar face, the
possibility that the observed effect might be the result of interference with general face
discrimination abilities, as opposed to self-recognition ought to be considered. Whilst the likelihood
of finding these results from such interference is very unlikely, future studies may wish to include
this control condition.
This finding improves our understanding of the self-other discrimination process but future TMS
research on empathy ought to expand on this to explore more directly the link between this process
and the observation of another individuals emotional state. Given that the EBA and the parietal
cortex both seem to play a role in self-other discrimination, further research may benefit from
examining how they activate independently during social interactions or connectedly.
There are several general limitations that should be taken into consideration when using TMS.
When using TMS-induced MEPs as a research method it should be remembered that similar
electrophysiological output can result from different neural stimulation, and also that these
measurements are not based on the activity of single neurons but rather on the mass activation of
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neural pathways. These limitations make it difficult to infer whether shared neural activations
indicate shared emotional representations. Future TMS research ought to be used alongside more
precise techniques such as multivariate pattern analysis, which has been used to show that the same
local activation patters are used when both experiencing pain and observing others (CorradiDellAcqua, Hofstetter, & Vuilleumier, 2011).
A related limitation is the correlational nature of MEP-TMS (Logothetis, 2008), which only shows
the experience of empathy coinciding with neural reactivity. To draw further causal conclusions
about shared activations being a necessary condition for empathy, these methods ought to be used
alongside neurostimulation or lesion studies. Neurostimulation is also limited by the unavailability
at this time of any methods that would allow the noninvasive stimulation of regions such as the
insular and cingulate cortices, which have both been identified as being important for mentalizing
(Fan, Duncan, de Greck, & Northoff, 2011), as they are deep under the cerebral mantle. Deeper
TMS techniques are required for a thorough examination of the involvement of these regions. It
should also be noted that each of the components that has been examined is on its own insufficient
to account for empathy. Future studies ought to take into account the role that each of them plays
and study them alongside one another.
The majority of TMS studies examining empathy have only looked at the negative effect on its
underlying components. However it is worth mentioning, briefly, the research examining the
possible application of TMS in order to enhance empathy in clinical populations where it may be
deficient. A small number of studies have shown that it may be possible to use TMS to influence
brain activity to enhance the levels of empathy experienced by individuals. Working with adults
with autism spectrum disorder (ASD), Theoret et al. (2005) first showed that TMS may be used to
increase motor resonance reactivity in populations with neuroatypical levels of empathy. Whilst
impairments usually result from low frequency rTMS, high frequency rTMS causes an increase in
excitation, and may be used as a method of treating deficiencies in empathy. One case study
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(Enticott, Kennedy, Zangen, Fitzgerald, 2011) reported that a woman with ASD, following nine
sessions of high frequency rTMS to the bilateral medial prefrontal cortex, had reported higher levels
of social functioning (measured using the IRI, AQ, and Ritvo Autism-Asperger Diagnostic Scale) as
well as feeling an increased capacity in empathy and perspective taking which her family
subsequently corroborated. However, when considering the use of TMS as a form of intervention, it
is important to make certain ethical considerations. Notably, many of the clinical conditions where
TMS may be beneficial are neurodevelopmental disorders meaning young children could benefit
from such interventions. However, little research has been conducted into the safety of TMS in
children (Rossi, 2011) and this is something which should be explored further before popularising
this intervention technique.
To conclude, studies using TMS have advanced out understanding of the processes underlying
empathy. They have shown that observing pain in others provokes the same neural response as if the
observer themselves were experiencing pain. They have shown that this resonance process may be
modulated by top-down mentalising such as racial bias. They have also provided the foundations
for further research into the process of attributing the source of internal affective states to the self or
other. The current literature indicates that TMS is a promising technique to use in understanding
empathy and may provide the area of empathy research with unique contributions, given its
exclusive ability, unlike other imaging techniques, to modulate the neural processes involved in the
different components of this system. Whilst imaging methods cannot examine causality, they have
provided much of the information which has lead to further research with TMS. Furthermore, it
would be simplistic to reduce empathetic experience neural activity alone such as cortical
excitability is a sufficient accurate indicator for empathy. It is therefore important to include
subjective accounts and behavioural measures of empathy alongside TMS, as well as using it in
conjunction with other brain imaging methods. Fortunately, the use of TMS to study empathy and
its underlying processes is becoming increasingly popular (Hetu et al., 2012).
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