!"##$#%&'&()*&%+,-%&"*.

/+("$,0%+%,)$1
)*2"#"3"&(%",()#2#)(+("$,

!"#$%&'(#)*+

!"#$%&#$%'%()*+$,*-"#*.%($/-/0#
12/#$2#3
!
"##$!%&'()**&+
!
,-./01!203/!#45
67"!%89%88*:;%%8+*)8%<)+<&=)=

1 23
Your article is protected by copyright and
all rights are held exclusively by Springer
Science+Business Media B.V.. This e-offprint
is for personal use only and shall not be self-
archived in electronic repositories. If you
wish to self-archive your work, please use the
accepted author’s version for posting to your
own website or your institution’s repository.
You may further deposit the accepted author’s
version on a funder’s repository at a funder’s
request, provided it is not made publicly
available until 12 months after publication.

1 23
 Author's personal copy
Phenom Cogn Sci
DOI 10.1007/s11097-012-9253-3

Mirror systems and simulation: a neo-empiricist

interpretation

John Michael

# Springer Science+Business Media B.V. 2012

Abstract It is often claimed that the discovery of mirror neurons supports simulation
theory (ST). There has been much controversy about this, however, as there are
various competing models of the functional contribution of mirror systems, only
some of which characterize mirroring as simulation in the sense required by ST. But a
brief review of these models reveals that they all include simulation in some sense. In
this paper, I propose that the broader conception of simulation articulated by neo-
empiricist theories of concepts can subsume the more specific conceptions of simu-
lation presented by ST and by these other models, thereby offering a framework in
which each of these models may play a role. According to neo-empiricism, conceptual
thought in general involves simulation in the sense that it is grounded in sensory, motor,
and other embodied systems (Barsalou, Behavioral and Brain Sciences, 22, 577–609,
1999, Philosophical Transactions of the Royal Society of London: Biological Scien-
ces, 364, 1281–1289, 2009; Barsalou et al., Trends in Cognitive Sciences, 7(2), 84–
91, 2003; Prinz 2002, Mind & Language, 25(5), 612–621, 2010; Glenberg and
Robertson, Journal of Memory and Language, 43, 379–401, 2000). Crucially, the
term “simulation” here refers not to simulations of a target agent’s experience in the
sense endorsed by simulation theory but to the activation of sensory, motor, affective,
and introspective representations. This difference does not entail that neo-empiricism
must be in competition with ST—indeed, I will propose that ST can be embedded as a
special case within neo-empiricism.

Keywords Neo-empiricism . Mirror neurons . Simulation theory . Embodied

cognition . Theory of mind . Mindreading

J. Michael (*)
Aarhus University, Tuborgvej 164, 2400 Copenhagen, Denmark
e-mail: joal@dpu.dk
 Author's personal copy
J. Michael

Introduction

Mirror neurons, which were originally discovered in macaque monkeys using single-
cell recordings, are neurons that are active when an animal is either executing a
particular action or observing another agent executing the same or a similar action.
Since it is only rare that single-cell activity can be recorded in human subjects (but
see Mukamel et al. 2010), mirror neuron research in humans has depended largely on
techniques for measuring (fMRI, EEG, MEG) or modulating (TMS) activation in
brain regions rather than in individual neurons. Thus, many researchers prefer to
speak of a “mirror system” or “mirror neuron system,” of which a brain region may
be considered a part if it is activated during the execution of an action as well as
during the observation of the same action (Frith and Singer 2008). Going by this
broader definition,1 the mirror system in humans can be localized to the ventral
premotor cortex (BA44/6), inferior parietal lobule, and somatosensory areas (BA2;
Buccino et al. 2001; Iacoboni et al. 2005; Rizzolatti and Craighero 2004). Mirror
systems have also been found for affective states and tactile sensations (Keysers et al.
2004; Singer et al. 2004; Adolphs 2003; Bastiaansen et al. 2009).
Research into mirror systems has attracted the interest of philosophers involved in
the mind reading debate,2 largely because it appears to corroborate the basic picture
of social cognition offered by simulation theory as an alternative to theory theory.
According to theory theory, social cognition is enabled by the ascription of unob-
servable mental states, which are defined in terms of their nomological relations to
perception, to behavior, and to other mental states (Carruthers 2009; Gopnik 1993;
Baron-Cohen 1995; Leslie 2000). Simulation theory, in contrast, is based on the idea
that we can understand others by “putting ourselves in their shoes,” or taking their
perspective, and that the representation of nomological psychological relations is
therefore superfluous. The common denominator of the various versions of simula-
tion theory is that predicting and/or understanding others’ actions and/or emotions
involves undergoing (simulating) the same procedures that we would undergo if we
ourselves were deciding upon, planning, or executing an action in the same circum-
stances or experiencing the same emotion (Gordon 1995, 2007; Goldman 1993, 2006,
2008; Heal 1986). Hence, it has been claimed that simulation theory predicts the kind
of first-/third-person overlap that mirror systems appear to constitute (Gallese and
Goldman 1998).
Apart from some fundamental concerns about whether mirroring really occurs at
all (e.g., Dinstein et al. 2007; Hickok 2009), there has also been a great deal of
controversy about whether mirror systems instantiate the kind of simulation that ST
predicts. Indeed, there are several distinct models of the functional contribution of
mirror systems, only some of which characterize mirroring as simulation in the sense
of ST. Interestingly, however, most of these models do involve some kind of simu-
lation. My aim here will be to show how the broader conception of simulation
articulated by neo-empiricist theories of concepts can subsume the more specific
1
I will use the term “mirror system” rather than “mirror neuron,” except where the context demands
otherwise.
2
I will use the term “mind reading” to refer to our perfectly ordinary ability to understand others as having
mental states of various sorts, including emotions, sensations, beliefs, and desires, as well as the sorts of
predictions and explanations of their behavior that such an understanding makes possible for us.
 Author's personal copy
Mirror systems and simulation: a neo-empiricist interpretation

conceptions of simulation presented by ST and by each of these models, and thereby

integrate these various models within a single theoretical framework.
According to neo-empiricism, conceptual thought in general involves simulation in
the sense that it is grounded in sensory, motor, and other embodied systems (Barsalou
1999, 2009; Barsalou et al. 2003; Prinz 2002, 2010; Glenberg and Robertson 2000).
The basic idea is that by perceiving and/or acting upon objects of a particular type, a
neural network for representing objects of that type is formed. When one subsequent-
ly perceives objects of that type, this network becomes active, presumably because
objects of that type have a set of features (lines, shapes, colors, sounds, contextual
factors, etc.) roughly in common and/or afford a roughly common set of actions. This
overlap in one’s responses is what enables one to categorize the object and, thus, also
to draw inferences about properties the objects are likely to have on the basis of their
membership in that category. Indeed, neo-empiricists maintain that imagining and/or
reasoning about objects that one is not presently encountering is also subserved by the
activation of such neural networks, thereby simulating the experience of perceiving or
acting upon the object(s) in question. Crucially, though, what is simulated is a past or
possible experience (or pattern of activation) of one’s own—not the experience of
some other target agent, as envisioned by ST.
The first part of this paper (see “Mirror systems: necessary but not sufficient” to
“What kind of simulation do mirror systems instantiate?”) will articulate and assess
the thesis that the discovery of mirror systems provides empirical support for ST. This
thesis can be analyzed into two claims:
(a) Mirror systems are involved in understanding others’ intentions or emotions.
(b) The way in which they are involved supports ST.
I will be giving qualified support to both (a) and (b). Starting (“Mirror systems:
necessary but not sufficient”) with claim (a), I will present theoretical and empirical
points in support of the view that mirror systems play a substantial role and are
perhaps necessary, although not sufficient, for understanding at least some intentions
or emotions. Although ST can easily accommodate this point, it does so by adding
non-simulative elements. As I will explain further below, neo-empiricism accommodates
the same point by adding further simulative elements. This difference does not constitute
an advantage for neo-empiricism over ST, but it will be an important point to
bear in mind in reflecting on the relationship between ST and neo-empiricism
and on the consequences of applying the neo-empiricist conception of simulation to the
interpretation of mirror systems.
Turning to (b), I will (“What kind of simulation do mirror systems instantiate?”)
review four leading models of the functional contribution of mirror systems to social
cognition. Although they differ in important respects, it turns out that each of them
involves a kind of simulation. The crucial point, however, is that only one of them
involves the kind of simulation envisioned by simulation theory, whereas all four
instantiate the kind of simulation postulated by neo-empiricism.
The next step (“Neo-empiricism: the other simulationist program”) will be to
characterize neo-empiricism in some detail and formulate the central predictions that
it generates about the role of the motor system (and other embodied representations)
in social cognition. Apart from being able to subsume and integrate all four of the
models reviewed here, neo-empiricism has the additional advantage of embedding
 Author's personal copy
J. Michael

mirror neuron research in a broader empirical context, which enables it to draw

support from a greater body of empirical research into the contribution of motor
and other embodied systems in cognition in general, thus increasing its base of
empirical support, its explanatory scope, and its theoretical coherency.
The final section (“Neo-empiricism and social cognition: some general issues”)
offers some general reflections about the relationship between ST and neo-empiricism
and considers the option of embedding ST as a special case within neo-empiricism. It
concludes with a brief discussion of the significance of neo-empiricism for debates
about embodied social cognition.

Mirror systems: necessary but not sufficient

It is often asserted that simulation theory predicts the kind of first-/third-person

overlap, or matching relation, that mirror systems appear to constitute (Gallese and
Goldman 1998; Goldman 2006, 2008). As noted in the “Introduction,” this assertion
can be analyzed into two claims:
(a) Mirror systems are involved in understanding others’ intentions or emotions.
(b) The way in which they do so supports simulation theory.
In this section, I will be assessing claim (a); in the next section, I will move to
claim (b). There are theoretical and empirical reasons to be wary of far-reaching
claims about the contribution of mirror systems to understanding intentions and
emotions. Let me start with two theoretical points. First, action understanding
appears to require a more abstract kind of representation than motor representation
since one action can be carried out with different movements and different actions can
be carried out with one and the same movement in different contexts. Proponents of
robust interpretations of mirror systems have addressed this issue. In an fMRI study,
Iacoboni et al. (2005) found differential activation in premotor areas in subjects
observing video clips of a hand performing the same movement in different contexts.
The authors infer that this differential activation reflects the integration of contextual
information for action interpretation. But, as Jacob (2008) points out, the differential
activation may result from the integration of contextual information occurring else-
where in the brain without contributing to it. Secondly, understanding someone’s
intention or emotion involves ascription of that intention or that emotion—simply
mirroring someone does not count as understanding that a particular state refers to
them rather than to oneself. It therefore seems unlikely that direct matching could be
sufficient for understanding intentions or affective experiences. But it may still be
necessary, i.e., in combination with other resources (e.g., contextual information,
concepts) and/or processes (e.g., ascription).
Given, then, that mirror system activation is not likely to be sufficient for under-
standing intentions and/or emotions, might it be necessary? There is some support for
an affirmative answer. Some transcranial magnetic simulation (TMS) studies have
found that inhibiting areas of the motor system impairs social cognitive skills. TMS-
induced inhibition of Broca’s area, for example, causes subjects to have difficulties
imitating observed actions, even if they do not have difficulties performing the same
actions in non-imitative scenarios (Heiser et al. 2003). Applying TMS to the inferior
 Author's personal copy
Mirror systems and simulation: a neo-empiricist interpretation

frontal gyrus has also been shown to impair subjects’ ability to estimate the weight of an
object lifted by a person in a video, but not their ability to estimate the weight of a bouncing
ball or the duration of a video (Pobric and Hamilton 2006). In the affective domain, the
evidence is even stronger. Wicker et al. (2003) found an overlap in activation between
scenarios where subjects experienced foul odors and when they saw others sniffing
the same foul odors. That this overlap is essential for understanding that the target
person is experiencing disgust is suggested by paired deficits in experience and
understanding of disgust (Adolphs 2003). Similar findings have been reported for
pain and fear (Singer et al. 2004; Adolphs et al. 2005; for a review, see Bastiaansen et
al. 2009). There is also evidence that mirroring others’ emotional facial expressions
plays a causal role in recognition of their emotions (Oberman et al. 2007).
In sum, although no final conclusions can yet be reached, there is presently good
reason to believe that mirror neuron activation is necessary, although not sufficient,
for understanding at least some intentions and emotions. Thus, we can provisionally
endorse claim (a) about the contribution of mirror systems to social cognition. What
does this mean for ST? It is important to note that none of the leading versions of ST
merely asserts such a matching relation and leaves it at that. In order to account for
the contribution of a simulation (or “mirroring event”) to mind reading, simulation
theorists all indeed make room for concepts (although they differ about whether
mental concepts are prerequisites to simulation) and also include an account of
ascription, i.e., how the simulated mental processes are separated from one’s own
mental processes and understood to refer to the target agent.
In fact, Goldman has also argued that, apart from the issue of ascription, it is also
necessary for ST to give an account of how the mental state that is the output of a
simulation is identified and categorized (Goldman 2006, p. 259). This is no trivial
challenge, however, since Goldman rejects the functionalist account of mental con-
cepts endorsed by theory theory (TT), i.e., he denies that mental states are differen-
tiated on the basis of representations of the nomological relations that theory theorists
believe are used to derive predictions about behavior from constellations of mental
states. The solution that Goldman has consistently advocated, albeit with very
significant modifications over the years, is introspectionism (Goldman 1993, 2006).
In the 1990s, he maintained that types of mental states are identified and categorized
on the basis of phenomenal properties (Goldman 1993). Goldman now suggests that
it is not phenomenal but neural properties that are introspected and has developed a
theory of introspection as unconscious monitoring (Goldman 2006). A proper assess-
ment of this proposal is beyond the scope of this paper,3 but it is worth noting that at
least one leading version of ST is committed to an introspectionist account of how
mental states are identified and categorized in order for simulations to contribute to
mind reading.
Thus, it is not a problem for ST that mirroring—or any other kind of simulation—
is likely to be only one component among others in a larger network underlying any
particular instance of mind reading. Nevertheless, there are two points to note about
what simulation theorists add to the mix in addition to the simulation that is at the
core of the theory. The first is that the further elements which they add are themselves
non-simulative—i.e., they do not instantiate simulation in the perspective-taking sense

3
But see Hill (2007).
 Author's personal copy
J. Michael

endorsed by ST. Secondly, at least one (highly influential) version of ST is committed to

giving an account of how mental states are differentiated on the basis of internally
accessible criteria. Again, these two points do not undermine ST, but they will be
relevant in discussing the relation between ST and neo-empiricism further below.
Having provisionally endorsed claim (a) and considered the consequences for
simulation theory, let us turn to claim (b). Does the role of mirror systems in social
cognition support simulation theory? I will now briefly review four models of the
functional contribution of mirror systems, each of which involves simulation, but
only one of which involves the kind of simulation envisioned by simulation theorists.

What kind of simulation do mirror systems instantiate?

Direct matching

The most straightforward and also most robust way of articulating the link between
mirror systems and simulation is the so-called direct matching hypothesis, according
to which the standard means of understanding others’ emotions or intentions is by
producing a response in the mirror system that matches the target state:
…the fundamental mechanism that allows us a direct experiential grasp of the
mind of others is not conceptual reasoning but direct simulation of the observed
events through the mirror mechanism. (Gallese et al. 2004)
Apart from the question of whether this sort of matching relation is likely to be
sufficient for understanding intentions or emotions or other mental states, one must
also ask whether it corresponds at all to the functional contribution that mirror
systems make. Surely this is suggested by the term “mirror neuron,” but it turns out
not to be entirely clear that this is the case.
Consider an argument formulated by Csibra (2008). He points out that only a
subset of mirror neurons is strictly congruent. Strictly congruent mirror neurons fire
when observing or performing one and the same action (same type of grasp and same
object). Many other mirror neurons are responsive to multiple actions. Beyond this,
many mirror neurons fire when one action is executed or when a functionally related
action is observed. Taken together, they constitute the class of “broadly congruent”
mirror neurons. Altogether, broadly congruent mirror neurons make up something
like 60% or 70% of all mirror neurons (Fogassi and Gallese 2002). The upshot of
Csibra’s criticism here is that only the strictly congruent mirror neurons would
successfully match an observed action with the activity patterns that are present when
the same action is executed. Therefore, if mirror neurons have anything to do with
understanding others’ intention, it is—at least for most mirror neurons—not by direct
matching.

Inverse modeling

Some theorists, such as Csibra himself (Csibra 2008; Jacob 2008, 2009), conclude
that mirror neurons do not play a role in identifying or ascribing intentions but
perhaps in predicting the ongoing motor realization of prior intentions, which are
 Author's personal copy
Mirror systems and simulation: a neo-empiricist interpretation

ascribed by other means. Jacob (2008) writes: “the job of MN activity is to compute
motor commands from a representation of the agent’s prior intention, not to represent
the agent’s prior intention….” This proposal assumes that ascription occurs more or
less in the way envisioned by theory theory, but in fact also contains a broadly
simulationist idea, namely that the observer’s own decision-making and action-planning
resources are employed when she seeks to predict the agent’s behavior. The simulation
in question is not a case of perspective taking, though, but of inverse modeling in the
sense common among computational theorists of action (e.g., Wolpert 1997; Wolpert et
al. 2003). Inverse models take a representation of a desired state of affairs (or prior
intention) as input and calculate an appropriate motor command for achieving that
goal, i.e., inferring backward from a goal state to the motoric means of achieving that
goal state. In contrast to the direct matching view, then, Csibra and Jacob do not grant
mirror neurons any role in understanding intentions. For them, the functional role of
mirror neurons is not to provide an alternative to conceptual understanding but indeed
depends on prior conceptual processing and ascription of an intention via the
application of mental concepts.
The model appears to be supported by the observation that most mirror neurons are
congruent in a broad rather than a narrow sense since it claims that mirror neurons do
not match but predict and would therefore represent not an observed movement but a
movement closely related to the observed movement. But it is crucially important to
know just what those broadly congruent mirror neurons are doing if not mirroring in a
narrow sense. That is, does their activation correspond to activation that would be
present in the observer’s brain if she were carrying out an action likely to follow upon
the presently observed action? This may be the case, but there are other possibilities
as well—e.g., the activation during observation may correspond to the activation that
would be present if the observer were carrying out a complementary response. And as
it happens, there are in fact data that support this alternative possibility.

Predictive coding

The predictive coding model, put forth by Kilner et al. (2007), is an application of the
predictive coding framework which is based on the conception of a hierarchy of
reciprocally connected models. Each model generates predictions about the repre-
sentations at the immediately subordinate level. These predictions are compared with
the actual state of the subordinate-level model, and a prediction error is returned to the
higher-level model, which is revised and then generates a new prediction. By this
process, the interconnected models are continuously updated and the prediction
errors minimized. Mirror systems, according to this proposal, would have the
function of generating relatively low-level, i.e., kinematic models of observed
behavior. Simultaneously, more abstract models would be generated at higher
levels of a hierarchy. Hence, a relatively high-level model yields a prediction
about an observed agent’s motor commands, while an intermediate model in
turn yields a prediction about the kinematics resulting from that motor com-
mand. This prediction is compared with the representations at a lower level, at
which mirror systems predict the kinematics of the observed movement. The
comparison of this prediction with the actual observed movement generates a
prediction error, which is used to revise our representation of the agent’s motor
 Author's personal copy
J. Michael

command in order to minimize the prediction error between the observed and inferred
kinematics. Similarly, our representation of the agent’s goal is revised to minimize the
prediction error between the predicted and inferred motor command.
The crucial difference between Csibra and Jacob’s inverse modeling model and the
predictive coding model is that the former is unidirectional whereas the latter is
bidirectional. According to Csibra and Jacob, mirror neuron activation depends upon
prior ascription of an intention and is not itself an input to mind reading. For
the predictive coding model, on the other hand, mirror neuron activation is not
only modulated by higher-level representations but also reciprocally modulates
higher-level representations. Hence, the predictive coding model—but not Csibra and
Jacob’s inverse modeling model—allows that activation in mirror systems can
contribute to action interpretation.
The functioning of mirror systems in this model, as in the inverse modeling model,
involves simulation in the sense of using one’s own motor system to dynamically
model a target agent’s future actions. But, again, this kind of simulation need not be
characterized as perspective taking (and the proponents of these models do not do so).
Moreover, both models integrate mirror systems with other processes (e.g., process-
ing perceptual information, integrating contextual and background information,
drawing theoretical inferences) that do not lend themselves at all to characterization
in terms of simulation in the sense of perspective taking.

Response modeling

For example, Newman-Norlund et al. (2007) found that the “human mirror neuron
system” (specifically, right inferior frontal gyrus and bilateral inferior parietal lobes)
is more active when observers are simultaneously preparing a complementary action
than when they are preparing an imitative action. They take this finding to suggest
that the function of these neurons lies in “dynamically coupling action observation to
action execution.” Note that, if this is the case, they would not be simulating the
observed agent’s movement or intention in the sense of simulation theory, but would
nevertheless be important for social cognition. Would they still be simulating in any
interesting sense?
In fact, there is a related usage of the term simulation which would apply quite well
here. The idea is that mirror systems could enable the interpreter to simulate the
experience of performing some other action which is related to the observed action or
some other emotional state which is related to the observed emotional state, for
example, an appropriate response. In this case, one would still be simulating an
experience. I would suggest that this sort of simulation may still be used in under-
standing the other person’s intention or emotion. The motor system may initiate some
complementary responses to observed behaviors on the basis of a fairly low level of
information processing. This sort of response would help maintain social coordina-
tion. More frontal areas such as vmPFC and rTPJ, which constitute the mentalizing
network (Castelli et al. 2002), would subsequently process not only the observed
behavior but also the interaction, of which the agent’s own complementary response
is a part. If the motor system is inhibited before the complementary response is
performed, the activation could be regarded as a simulation of the complementary
response and could nevertheless be an input to the mentalizing network. Clearly, the
 Author's personal copy
Mirror systems and simulation: a neo-empiricist interpretation

kind of simulation at issue in this sort of case is different from what simulation
theorists have in mind when speaking of simulation as a means of perspective taking.
But it is perfectly compatible with the neo-empiricist conception of simulation.
In fact, there are other studies that fit well with this interpretation and also use
simulation in this sense. For example, Iacoboni (2005) (pp. 127–128) reports a study
of mothers’ responses to pictures of babies expressing emotions. Unsurprisingly,
there were stronger responses in mirror neuron areas and also in emotional brain
centers (insula and amygdala) when looking at pictures of their own babies as
opposed to other babies. What is interesting is that there was also increased activation
in the pre-supplementary motor area (pre-SMA), which is anatomically linked closely
with mirror neuron areas and seems to control and modulate mirror neuron activation.
The pre-SMA is an “important region for complex motor planning and motor
sequencing—that is, for putting together a series of concatenated actions” (Iacoboni
2005, p. 128)—i.e., for actions more complex than the mere facial expressions they
are observing and mirroring. Iacoboni’s gloss is that the mothers’ mirror systems are
(motorically and emotionally) mirroring and that this mirroring activity additionally
“triggers a whole cascade of other automatic simulative brain responses,” which may
in turn increase mirror system activation further (Iacoboni 2005, p. 129). Note that
Iacoboni uses the term “simulative” for this response selection process in a broader
sense that is compatible with neo-empiricism, but not with simulation theory, i.e., it is
not the other person’s perspective that is being modeled but one’s own possible
response.

Comparing the models

It is important to point out that none of the models discussed so far intrinsically
excludes the possibility that they all might be correct, i.e., that neural circuits with
mirroring properties might instantiate direct matching, inverse modeling, and response
selection. Indeed, the observation that some mirror neurons exhibit strict congruence
while others exhibit broad congruence suggests that they may not all serve the same
function. A natural interpretation is that strictly congruent mirror neurons instantiate
either direct matching or predictive coding, whereas broadly congruent mirror neu-
rons instantiate inverse modeling or response modeling. Moreover, many of the
studies discussed so far lend themselves to a pluralist interpretation. Recalling the
results of Newman-Norlund et al. (2007) mentioned above, one might argue that they
in fact support the view that mirror systems have multiple functions. After all,
performing a complementary action could involve direct matching, inverse modeling,
and/or predictive coding, plus the additional task of responding in a complementary
fashion, so it is no wonder that there is more activation in the complementary action
scenario than in the scenario where there is only imitation, which involves minimal
additional burdens beyond mirroring. The results, properly considered, suggest that
mirror systems have a response-selective function in addition to direct matching,
predictive coding, and/or inverse modeling.
Given the possibility that mirror systems might instantiate various types of model,
an interpretation that subsumes multiple models would be in a strong position in that
it would be able to claim support from all the data drawn upon in favor of any one of
the models. In the next section, I will be arguing that neo-empiricist theories of
 Author's personal copy
J. Michael

concepts offer the possibility of developing just such a “supermodel.” Neo-

empiricism articulates a broad conception of simulation that can subsume the more
specific conceptions of simulation presented by ST and by each of these models,
thereby offering a framework in which each of these models may play a role.

Neo-empiricism: the other simulationist program

Neo-empiricism

I would like to begin with a brief sketch of neo-empiricist theories of concepts. The
basic idea is that conceptual thought, rather than taking place in an amodal symbolic
code (i.e., a “language of thought”), involves the same modality-specific neural
activity as perception. Let me explain how this is supposed to work with a simple
example borrowed from Barsalou (1999). When one sees a car, neural feature
detectors are active in the visual system. Conjunctive neurons in a nearby area
conjoin the active features and store them in memory. This is referred to as the
“capture procedure.” These sets of conjunctive neurons also account for the trans-
modal nature of concepts, namely by integrating the feature detection activity that
occurs during visual perception of the car with feature detection activity that is active
in other modality-specific systems, such as the auditory system.
With the help of selective attention, the capture procedure can be restricted to
components of scenes (e.g., a car on the street) rather than entire scenes (e.g., a street).
Thus, when focusing attention on the car, specific (modal and conjunctive) neurons
become active through the capture procedure. Memory integration causes this pattern of
neural activation to be integrated with similar previous patterns. As a result of such
cumulative integration, a network for representing cars is formed. When, in the future, a
subsequent car is perceived, or when one reasons about cars, this network becomes
active, presumably because cars have a set of features (lines, shapes, colors, sounds,
contextual factors, etc.) roughly in common, and the perception of these features causes
the activation of common neurons, thereby simulating the sensory perception of the car.
Since simulations, on a neo-empiricist view, instantiate concepts, their role is to
enable the performance of conceptual tasks, such as drawing inferences. If one then
perceives, for example, the sound of the car, a simulator will be activated that also
includes visual representations associated with cars. These visual representations
encode additional information concerning the car, such as the property of having
four wheels, and thus enable one to infer that the object producing the sound in
question will have four wheels.
Crucially, this notion of simulation differs from that of simulation theory insofar as
it does not apply uniquely to taking the perspective of (0simulating) a different
person. Rather, it is a feature of conceptual thought in general. As Barsalou puts it,
“simulation constitutes a central form of computation throughout diverse forms of
cognition, where simulation is the reenactment of perceptual, motor and introspective
states acquired during experience with the world, body and mind” (Barsalou 2009).
Barsalou indeed calls the networks that coordinate the reactivation of multimodal
representations “simulators.” It is worth noting that Prinz (2002, 2010), who endorses
a similar theory, also uses the term “simulation” in this sense.
 Author's personal copy
Mirror systems and simulation: a neo-empiricist interpretation

It is worth emphasizing that simulators instantiate a plurality of representations in a

dynamic fashion, i.e., in temporally extended sequences. That is, simulators include
not only perceptual representations but also motor representations, as well as affective
states and what Barsalou (1999) calls “introspection,” which he employs in a neutral
sense to refer to various kinds of self-directed (conscious or unconscious) cognitive
processes, such as monitoring one’s own affective states and action plans and making
metacognitive judgments. The plurality and dynamic nature of the representations
integrated in simulators is central to neo-empiricist accounts of complex and abstract
concepts, which do not easily lend themselves to reduction to an exclusively senso-
rimotor basis. A relatively complex concept, such as “anger,” for example, may
involve several components, such as arousal and neural activation typically present
when one is in an angry state, perceptual representations of angry behavior, and
perhaps also typical cognitive operations underlying judgments typically associated
with “anger” (e.g., the judgment that a given action has caused harm or was unfair,
the inference that revenge or punishment is appropriate, etc.). Thus, the component of
a concept that could plausibly be instantiated by a mirror system (i.e., an affective
mirror system), namely the arousal and/or neural activation associated with anger, is
only one part of a broader network.
Or, to take an example of a concept that could be instantiated in part in the motor
system, consider the concept of “drink.” This is a relatively abstract concept, insofar
as there are many things one could drink, receptacles one could drink from, motor
programs that could be engaged, etc. A simulator for this concept could plausibly
include visual representations of typical fluids and receptacles, as well as representa-
tions of people drinking. It could also include a haptic representation of fluid in one’s
mouth, an interoceptive representation of thirst, and various motor programs for
gasping a receptacle, for swallowing, etc. Moreover, these representations could be
integrated in a fashion that includes information about their typical sequences (e.g., a
representation of thirst, various visual and motor programs, then the extinguishing of
the thirst). Although none of these components would alone be sufficient to constitute
the concept of drinking, such a constellation of various kinds of representation linked
up by Hebbian learning may be.
In short, neo-empiricism predicts that one would find the kind of activation of
embodied systems that mirror systems appear to constitute, and it also predicts that
the contribution of these embodied systems would be contingent upon their integra-
tion with broader networks.

A neo-empiricist supermodel?

The plurality of the representations integrated in neo-empiricist accounts of concepts

provides a sound basis for developing the sort of “supermodel” of mirror systems
alluded to above (“Comparing the models”). If so, it would be in a strong position in
that it would be able to claim support from all the data drawn upon in favor of any one
of the models. Moreover, it could contribute additional theoretical coherency by
showing how the more specific models relate to each other. Thus, a neo-empiricist
supermodel could specify simulators in such a way that they include neural circuits
instantiating different sub-models. For example, it is plausible to suppose that a
simulator instantiating a particular action concept would include links to various
 Author's personal copy
J. Michael

motor areas instantiating various sub-models and would activate these links in a
manner sensitive to the task at hand. For example, the task of imitating an observed
action would likely engage visual areas as well as motor areas instantiating a direct
matching model. Detached interpretation of the same action, on the other hand, may
engage the same visual areas, but motor areas instantiating inverse modeling or
predictive coding rather than direct matching. Fluid coordination of complementary
behavior in a joint action, in contrast, may be more likely to engage response
selection circuits.
Although there may be a limit to how well embodied theories can account for
complex and abstract concepts—such as concepts of some complex intentions and
other mental states—the sort of combination of diverse representations that neo-
empiricism proposes may work for some of the mental concepts that are important
for social cognition. Moreover, by making room for so-called conjunctive neurons to
coordinate modal representations (as discussed above in “Direct matching”), neo-
empiricist approaches already embed modal representations in a hierarchical organi-
zation that includes representations of varying degrees of abstraction. It remains an
open question how mental concepts should be mapped onto such a hierarchy—the
main thrust of a neo-empiricist approach to social cognition is to predict that modal
representations play a significant role in the dynamic simulations that underlie mental
concepts, not to deny that hierarchically superior neural circuits also play important
coordinating roles.
Rather than merely noting that mirror systems cannot be sufficient for constituting
mental concepts, it is likely to be more fruitful to regard the relationship between
mirror systems and mental concepts as akin to the relationship between embodied
representations and abstract concepts. Work on abstract concepts within the neo-
empiricist framework demonstrates the benefits of thinking about the constraints
upon abstract concepts imposed by the need to ground them within sensory, motor,
and other embodied systems. Mental concepts must, for example, be encoded in such
a way that they can be used to identify instances of their referents in real-life
situations, draw inferences about behavior that are context-sensitive, and guide one’s
one behavior in a way that is sensitive to one’s own mental states. In short, although it
is an open question just how well one can account for mental concepts with motor,
sensory, and other embodied representations, it seems highly likely that such embod-
ied representations at least partially constitute mental concepts, and mirror systems
are likely to be part of the story.

Further evidence for the broader, neo-empiricist conception of simulation

A further virtue of the neo-empiricist framework is that it can appeal to a broad base
of psychological and neuroscientific evidence in support of its claim that cognition in
general involves simulation in the sense of activating embodied systems. Behavioral
studies have, for example, revealed that modality switching and perceptual similarity
slow verification of features as typical—e.g., people are faster at verifying “loud” for
“blender” after “rustling” for “leaves” than after “tart” for “cranberries” (Pecher et al.
2003) and faster at verifying “mane” for “pony” after verifying “mane” for “horse”
than after verifying “mane” for “lion” (Pecher et al. 2003). Similar studies have
targeted the causal contribution of the motor system in conceptual processing—e.g.,
 Author's personal copy
Mirror systems and simulation: a neo-empiricist interpretation

people are quicker to verify that “Open the drawer” is a sensible sentence when their
response involves a pulling motion as opposed to a pushing motion (Glenberg and
Kaschak 2003).
In another study, conducted by Vermeulen et al. (2008), participants held either
pictures or sounds in working memory while confirming the typicality of features
(e.g., Is a lemon yellow? Is a blender loud?). They found that visual working
memory load increased reaction times on visual features, but not auditory
features, and that auditory load increased reaction times on auditory features,
but not visual features. Such load effects should not be expected on an amodal
account of concepts, but are exactly what neo-empiricism predicts.
Looking at the evidence from neuroscience, one study (Chao et al. 1999) has found
an overlap in the activation in perceptual areas when reading the names of familiar
categories and when seeing pictures of objects from those categories. Another study
(Simmons et al. 2003) has shown that the pattern of sensory activation during a
linguistic task is correlated with the perceptible features of the objects named in the
linguistic task, e.g., questions about blenders cause activation in auditory areas. TMS
studies also provide evidence that the motor system contributes causally to concep-
tual processing. For example, using TMS to facilitate the activation of a part of motor
cortex that is active during leg motions specifically facilitates comprehension of
sentences describing actions involving legs, whereas stimulating motor areas that
are active during arm motions specifically facilitates comprehension of sentences
describing actions involving arms (Pulvermüller and Hauk 2005).
Given all this evidence that embodied representations contribute to various concep-
tual tasks, the most sensible interpretation appears to be that simulation in the broader,
neo-empiricist sense is a prevalent feature of cognition and that mirror systems are an
instance of this. In contrast, the simulation theorist’s strategy of isolating mirror systems
and pointing to them as evidence of a specific kind of simulation appears arbitrary. This
does not imply, however, that neo-empiricism and ST need to be in competition with
each other. In the next section, I will address this issue directly, taking a closer look at the
relationship between these two simulationist programs and considering the option of
embedding ST as a special case within neo-empiricism. I will also tie together some
loose ends and offer some reflections about the significance of neo-empiricism for
debates about embodied social cognition.

Neo-empiricism and social cognition: some general issues

Simulation or simulation?

Although neo-empiricism does not entail simulation theorists’ claim that social
cognition is unique in that it involves simulation of the other person’s perspective,
it does not by any means rule out the possibility that simulation in the sense of ST is a
special case of simulation in the broader, neo-empiricist sense. This may even be an
attractive option for simulation theorists. Goldman and Shanton (2010), for example,
have explicitly considered the relationship between ST and Barsalou’s broader
simulationist program. They suggest that embedding ST within a neo-empiricist
account like Barsalou’s may strengthen ST. According to them, if simulation is in
 Author's personal copy
J. Michael

fact a central feature of cognition in general, as neo-empiricism claims, it is all the

more likely to be a central feature of social cognition as well. They stop short of
endorsing neo-empiricism, though, noting that ST need not commit to what is, after
all, a more far-reaching theoretical program.
It seems clear, however, that one could just as easily turn this point around. Insofar as
ST postulates a more specific kind of simulation, namely perspective taking, it could be
viewed as the more ambitious program. At least with respect to mirror systems, it seems
that any evidence for ST would also count as evidence for neo-empiricism because
perspective-taking simulation instantiated by mirroring would be evidence for neo-
empiricism since it would corroborate the claim that the reactivation of embodied
(motor) representations is a prevalent feature of cognition. The reverse is not true—
neo-empiricism, but not ST, could gain support from evidence of simulation in the sense
implied by the response modeling conception or by other models that do not involve
perspective taking but do involve the reactivation of motor representations.
There are three important qualifications to be made here. First, although evidence
that mirror systems instantiate ST supports not only ST but also neo-empiricism, it
may support ST more specifically. And, as ST, as it were, goes further out on a limb to
venture a more specific hypothesis, it should also be seen to gain even more
credibility from evidence substantiating that hypothesis. Secondly, recall that both
ST and neo-empiricism predict that the contribution of mirror systems would be
contingent upon their integration with other systems. As I noted above (“Mirror
systems: necessary but not sufficient” and “Neo-empiricism”), the difference lies in
what elements they add to the mix. Specifically, simulation theorists add various
components to account for the ascription of mental states that are not themselves
simulative. Additionally, insofar as Goldman wants to avoid the TT view that mental
states are differentiated according to the nomological relations they enter into, his
version of ST is also committed to an introspectionist account of how mental states
are identified and categorized.
Neo-empiricism also predicts that other embodied (perceptual, affective, proprio-
ceptive, etc.) representations are activated beyond the motor system, but it character-
izes this further activation as more simulation, i.e., its broader conception of
simulation is applicable to the additional elements that it adds to the mix as well.
Thus, although the activation of motor representations in off-line social cognition is
predicted by both ST and neo-empiricism, they make different predictions about the
broader networks that these motor representations are integrated into. It is in principle
possible to find evidence for components underlying ascription or introspective
categorization that involve amodally formatted representations. Such evidence, if
found, could support ST, but not neo-empiricism.
The third qualification (and this is a generalization of the second point) is
that, although a case of perspective-taking simulation instantiated by mirroring
would also be evidence for neo-empiricism, this is not true of evidence for
simulations in general. Thus, evidence for high-level simulation that does not involve
the motor system may support ST, but not neo-empiricism, if it could also be established
that the representations instantiating the simulation were not embodied representations
as required by neo-empiricism but were, for example, in an amodal format.
There is one other important point to raise about the relationship between these
two simulationist programs. For ST, simulation is understood as an alternative to the
 Author's personal copy
Mirror systems and simulation: a neo-empiricist interpretation

representation of nomological relations among mental states (and to behavior

and perception), which—according to TT—underlie conceptual abilities such as
categorization and inference. Simulations in the neo-empiricist sense, however, are not
considered as alternatives to but as components of conceptual processing. Thus, neo-
empiricism avoids the problem of having to explain how we identify and categorize
simulated mental states in order to ascribe them to others because the simulation of a
particular mental state is already the application of the concept of that mental state.
For conceptual thought, according to neo-empiricism, involves the activation of a
plurality of embodied representations, some of which would likely be in the motor
system (e.g., mirror systems) and some of which would likely lie in other embodied
systems. Thus, neo-empiricist simulations can involve, for example, the activation of
sensory representations from prior experiences as a second-person interactant or a
third-person observer in social cognition. A consequence of this is that simulation in
the neo-empiricist sense can instantiate TT as well as ST, and thus provides an
intriguing option for those interested in developing either a hybrid of TT and ST or
in moving beyond the TT/ST dichotomy and reflecting on how the processes
emphasized by advocates from both camps may complement each other.

Neo-empiricism and embodied social cognition

Applying neo-empiricism to social cognition also makes it possible to build bridges

between embodied approaches to social cognition, on the one hand, and more
traditionally cognitivist approaches, on the other. Proponents of embodied approaches
often claim that embodied and interactive processes render conceptual processes or
mind reading superfluous (e.g., Gallagher and Zahavi 2008; Gallagher 2001, 2007,
2008; Hobson 2002; Reddy 2008; Hutto 2004; De Jaegher 2009; De Jaegher and Di
Paolo 2007; Fuchs and De Jaegher 2009; De Jaegher et al. 2010). Although neo-
empiricism obviously concurs in ascribing a central role to embodied interaction in
(social) cognition, it differs from these approaches in that it does not conceptualize
embodiment as an alternative to conceptual processes or to mind reading and does not
argue that embodied processes replace mind reading or conceptual processes. Rather,
it espouses the more moderate view that some conceptual processes may be (at least
partially) constituted by embodied processes.
One reason why this sort of moderate view should be considered appealing is that
it invites attempts to show how the embodied processes engaged in interactions make
the tasks of understanding others and interacting with them different than they would
be if we really were just disembodied cognizers. For example, it is likely that
individual social cognitive processes have been shaped by the need to monitor,
coordinate, and modulate embodied processes engaged in interactions. Thus, rather
than conceiving of embodiment as a replacement for the individual cognitive pro-
cesses envisioned by traditional approaches to social cognition, it could be possible to
improve those approaches by integrating embodied processes into their models.
To see how neo-empiricism can contribute to this effort, consider the suggestion
made in the discussion of response modeling above (“Inverse modeling”). It was
proposed there that the embodied responses to others’ actions or emotions that one
initiates on the basis of quick, automatic, and unconscious low-level processes could
be monitored by and thus provide information to higher-level information processing.
 Author's personal copy
J. Michael

If so, then a natural hypothesis for neo-empiricists to advance would be that dynamic
simulations (online or off-line) underlying judgments about others’ intentions or
emotions may recruit not only perceptual areas that would be active if one were
observing another person but also motor or affective areas that would be active if one
were responding to others. This is just one example intended to demonstrate how
traditional approaches to social cognition might benefit by integrating embodied
processes into their models and how neo-empiricism can assist in this effort.

Conclusion

Since neo-empiricism is a theory about the format in which concepts are represented,
it is not in direct competition with other leading theories of concepts (e.g., theory
theory, prototype theory, exemplar theory). Similarly, when applied to social cogni-
tion and thus to mental concepts, it is not in direct competition with other leading
theories in the mind reading debate. Simulators in the neo-empiricist sense could, for
example, instantiate knowledge of nomological psychological laws as postulated by
theory theory, or simulations in the more specific sense of simulation theory, or some
other psychological process(es) underlying social cognition.
My aim here has not been to argue that neo-empiricism is correct and simulation
theory (or theory theory) false. Rather, I have tried to show that neo-empiricism is well
placed to integrate various models of the functioning of mirror systems by subsuming
the distinct senses of simulation that they employ under a broader conception of
simulation. Apart from this, neo-empiricism has the additional virtue that it embeds
mirror neuron research in a broader empirical context, which enables it to draw support
from a greater body of empirical research into the contribution of motor and other
embodied systems in cognition in general, thus increasing its base of empirical support,
its explanatory scope, and its theoretical coherency. Moreover, since neo-empiricism
claims that simulation can instantiate conceptual thought rather than providing an
alternative to it, it offers an intriguing option for those interested in developing either
a hybrid of TT and ST or in moving beyond the TT/ST dichotomy. Finally, I have argued
that a neo-empiricist approach to the role of embodiment in social cognition presents a
viable, moderate alternative to more radical approaches that conceptualize embodiment
as a replacement for individual social cognitive processes.

Acknowledgments Many thanks to two anonymous reviewers, both of whose suggestions helped a great
deal in improving this paper.

References

Adolphs, R. (2003). Cognitive neuroscience of human social behaviour. National Review of Neuroscience,
4(3), 165–178.
Adolphs, R., Gosselin, F., Buchanan, T. W., Tranel, D., Schyns, P., & Damasio, A. R. (2005). A mechanism
for impaired fear recognition after amygdala damage. Nature, 433, 68–72.
Baron-Cohen, S. (1995). Mindblindness: An essay on autism and theory of mind. Cambridge, MA: MIT
Press.
 Author's personal copy
Mirror systems and simulation: a neo-empiricist interpretation

Barsalou, L. (1999). Perceptual symbol systems. The Behavioral and Brain Sciences, 22, 577–609.
Barsalou, L. (2009). Simulation, situated conceptualization, and prediction. Philosophical Transactions of
the Royal Society of London: Biological Sciences, 364, 1281–1289.
Barsalou, L., Kyle Simmons, W., Barbey, A., & Wilson, C. (2003). Grounding conceptual knowledge in
modality-specific systems. Trends in Cognitive Sciences, 7(2), 84–91.
Bastiaansen, J. A. C. J., Thioux, M., & Keysers, C. (2009). Evidence for mirror systems in emotions.
Philosophical Transactions of the Royal Society B, 364, 2391–2404.
Buccino, G., Binkofski, F., Fink, G. R., Fadiga, L., Fogassi, L., Gallese, V., Seits, R. J., Zilles, K.,
Rizzolatti, G., & Freund, H.-J. (2001). Action observation activates premotor and parietal areas in a
somatotopic manner: An fMRI study. European Journal of Neuroscience, 13, 400–404.
Carruthers, P. (2009). How we know our minds: The relationship between metacognition and mindreading.
The Behavioral and Brain Sciences, 32, 121–182.
Castelli, F., Frith, C., Happé, F., & Frith, U. (2002). Autism, Asperger syndrome and brain mechanisms for
the attribution of mental states to animated shapes. Brain, 125(8), 1839–1849.
Chao, L. L., Haxby, J. V., & Martin, A. (1999). Attribute-based neural substrates in posterior temporal
cortex for perceiving and knowing about objects. Nature Neuroscience, 2, 913–919.
Csibra, G. (2008). Action mirroring and action understanding: An alternative account. In P. Haggard, Y.
Rossetti, & M. Kawato (Eds.), Sensorimotor foundation of higher cognition: Attention and perfor-
mance (pp. 435–458). Oxford: Oxford University Press.
De Jaegher, H. (2009). Social understanding through direct perception? Yes, by interacting. Consciousness
and Cognition, 18(2009), 535–542.
De Jaegher, H., & Di Paolo, E. (2007). Participatory sense-making: An enactive approach to social
cognition. Phenomenology and the Cognitive Sciences, 6(4), 485–507.
De Jaegher, H., Di Paolo, E., & Gallagher, S. (2010). Can social interaction constitute social cognition?
Trends in Cognitive Sciences, 14(10), 441–447.
Dinstein, I., Hasson, U., Rubin, N., & Heeger, D. J. (2007). Brain areas selective for both observed and
executed movements. Journal of Neurophysiology, 98(3), 1415–1427.
Fogassi, L., & Gallese, V. (2002). The neural correlates of action understanding in non-human primates. In
M. I. Stamenov & V. Gallese (Eds.), Mirror neurons and the evolution of brain and language (pp. 13–
35). Amsterdam: John Benjamins.
Frith, C., & Singer, T. (2008). The role of social cognition in decision-making. Philosophical Transactions
of the Royal Society B, 363, 3875–3886.
Fuchs, T., & De Jaegher, H. (2009). Enactive intersubjectivity: Participatory sense-making and mutual
incorporation. Phenomenology and the Cognitive Sciences, 8, 465–486.
Gallagher, S. (2001). The practice of mind: Theory simulation or primary interaction? Journal of Con-
sciousness Studies, 8(5–7), 83–108.
Gallagher, S. (2007). Simulation trouble. Social Neuroscience, 2(3/4), 353–365.
Gallagher, S. (2008). Direct perception in the intersubjective context. Consciousness and Cognition, 17(2),
535–543.
Gallagher, S., & Zahavi, D. (2008). The phenomenological mind. London: Routledge.
Gallese, V., & Goldman, A. (1998). Mirror neurons and the simulation theory of mind-reading. Trends in
Cognitive Sciences, 2, 493–501.
Gallese, V., Keysers, C., & Rozzolatti, G. (2004). A unifying view of the basis of social cognition. Trends in
Cognitive Sciences, 8(9), 396–403.
Glenberg, A. M., & Kaschak, M. P. (2003). Grounding language in action. Psychonomic Bulletin Review, 9,
558–565.
Glenberg, A., & Robertson, D. (2000). Symbol grounding and meaning: A comparison of high-
dimensional and embodied theories of meaning. Journal of Memory and Language, 43, 379–
401.
Goldman, A. (1993). The psychology of folk psychology. In A. Goldman (Ed.), Readings in philosophy
and cognitive science (pp. 347–380). Cambridge, MA: MIT Press.
Goldman, A. (2006). Simulating minds: The philosophy, psychology, and neuroscience of mindreading.
Oxford: Oxford University Press.
Goldman, A. (2008). Mirroring, mindreading and simulation. In J. A. Pineda (Ed.), Mirror neuron system:
The role of mirroring processes in social cognition (pp. 311–330). New York: Humana.
Goldman, A., & Shanton, K. (2010). The case for simulation theory. In A. Leslie & T. German (Eds.),
Handbook of “theory of mind”. New York: Psychology Press.
Gopnik, A. (1993). The illusion of first-person knowledge of intentionality. The Behavioral and Brain
Sciences, 16, 1–14.
 Author's personal copy
J. Michael

Gordon, R. (1995). Simulation without introspection or inference from me to you. In T. Stone & M. Davies
(Eds.), Mental simulation: Evaluations and applications (pp. 53–67). Oxford: Blackwell.
Gordon, R. (2007). Ascent routines for propositional attitudes. Synthese, 159, 151–165.
Heal, J. (1986). Replication and functionalism. In J. Butterfield (Ed.), Language, mind and logic (pp. 135–
150). Cambridge: Cambridge University Press.
Heiser, M., Iacoboni, M., Maeda, F., Markus, J., & Maziotta, J. (2003). The essential role of Broca’s area in
imitation. European Journal of Neuroscience, 17, 1123–1128.
Hickok, G. (2009). Eight problems for the mirror neuron theory of action understanding in monkeys and
humans. Journal of Cognitive Neuroscience, 21(7), 1229–1243.
Hill, C. (2007). Goldman’s theory of introspection (manuscript).
Hobson, R. P. (2002). The cradle of thought. London: Macmillan.
Hutto, D. (2004). The limits of spectatorial folk psychology. Mind and Language, 19(5), 548–573.
Iacoboni, M., Molnar-Szakacs, I., Gallese, V., Buccino, G., Mazziotta, J. C., & Rizzolatti, G. (2005).
Grasping the intentions of others with one’s own mirror neuron system. PLoS Biology, 3, 529–535.
Jacob, P. (2008). What do mirror neurons contribute to human social cognition? Mind and Language, 23(2),
190–223.
Jacob, P. (2009). The tuning-fork model of human social cognition: A critique. Consciousness and
Cognition, 18(1), 229–243.
Keysers, C., Wicker, B., Gazzola, V., Anton, J. L., Fogassi, L., & Gallese, V. (2004). A touching sight: SII/
PV activation during the observation and experience of touch. Neuron, 42, 335–346.
Kilner, J., Friston, K., & Frith, C. (2007). Predictive coding: An account of the mirror neuron system.
Cognitive Processing, 8, 159–166.
Leslie, A. (2000). Theory of mind as a mechanism of selective attention. In M. Gazzaniga (Ed.), The new
cognitive neurosciences (pp. 1235–1247). Cambridge, MA: MIT Press.
Mukamel, R., Ekstrom, A. D., Kaplan, J., Iacoboni, M., & Fried, I. (2010). Single-neuron responses in
humans during execution and observation of actions. Current Biology, 20, 750–756.
Newman-Norlund, R., van Shie, H., van Zuijlen, A., & Beckering, H. (2007). The mirror system is
more active during complementary compared with imitative action. Nature Neuroscience, 10(7),
817–818.
Oberman, L., Winkielman, P., & Ramachandran, V. S. (2007). Face to face: Blocking facial mimicry can
selectively impair recognition of impair emotional expressions. Social Neuroscience, 2(3–4), 167–178.
Pecher, D., Zeelenberg, R., & Barsalou, L. W. (2003). Verifying properties from different modalities for
concepts produces switching costs. Psychological Science, 14, 119–124.
Pobric, G., & Hamilton, A. (2006). Action understanding requires the left inferior frontal cortex. Current
Biology, 16, 524–529.
Prinz, J. (2002). Furnishing the mind: Concepts and their perceptual basis. Cambridge, MA: MIT Press.
Prinz, J. (2010). Can concept empiricism forestall eliminativism? Mind & Language, 25(5), 612–621.
Pulvermüller, F., & Hauk, O. (2005). Functional links between motor and language systems. European
Journal of Neuroscience, 21, 793–797.
Reddy, V. (2008). How infants know minds. Cambridge, MA: Harvard University Press.
Rizzolatti, G., & Craighero, L. (2004). The mirror-neuron system. Annual Reviews of Neuroscience, 27,
169–192.
Simmons, W. K., Pecher, D., Hamann, S. B., Zeelenberg, R., & Barsalou, L. W. (2003). fMRI evidence for
modality-specific processing of conceptual knowledge on six modalities. Meeting of the Society for
Cognitive Neuroscience, New York, March 2003.
Singer, T., Seymour, B., O’Doherty, J., Kaube, H., Dolan, R. J., & Frith, C. (2004). Empathy for pain
involves the affective but not sensory components of pain. Science, 303, 1157–1162.
Vermeulen, N., Corneille, O., & Niedenthal, P. M. (2008). Sensory load incurs conceptual processing costs.
Cognition, 109, 287–294.
Wicker, B., Keysers, C., Plailly, J., Royet, J., Gallese, V., & Rizzolatti, G. (2003). Both of us disgusted in
my insula: The common neural basis of seeing and feeling disgust. Neuron, 40, 4–655.
Wolpert, D. M. (1997). Computational approaches to motor control. Trends in Cognitive Sciences, 1, 209–
216.
Wolpert, D. M., Doya, K., & Kawato, M. (2003). A unifying computational framework for motor control
and social interaction. Philosophical Transactions of the Royal Society, London B, 358, 593–602.