Exp Brain Res (2010) 200:223–237
DOI 10.1007/s00221-009-2002-3
R EV IE W
Mirror neurons: from discovery to autism
Giacomo Rizzolatti · Maddalena Fabbri-Destro
Received: 12 June 2009 / Accepted: 27 August 2009 / Published online: 18 September 2009
© Springer-Verlag 2009
How the things started
In the winter of 1991 I (GR) sent to Nature a report on a
surprising set of neurons that we (Giuseppe Di Pellegrino,
Luciano Fadiga, Leonardo Fogassi, Vittorio Gallese) had
found in the ventral premotor cortex of the monkey. The
fundamental characteristic of these neurons was that they
discharged both when the monkey performed a certain
motor act (e.g., grasping an object) and when it observed
another individual (monkey or human) performing that or a
similar motor act (Di Pellegrino et al. 1992). These neurons
are now known as mirror neurons (Fig. 1).
Nature rejected our paper for its “lack of general inter-
est” and suggested publication in a specialized journal. At
this point I called Prof. Otto Creutzfeld, the then Coordinat-
ing Editor of Experimental Brain Research. I told him that I
thought we found something really interesting and asked
him to read our manuscript before sending it to the referees.
After a few days he called me back saying that indeed our
Wndings were, according to him, of extraordinary interest.
G. Rizzolatti (&) · M. Fabbri-Destro
Dipartimento di Neuroscienze, Sezione Fisiologia,
Università di Parma, via Volturno, 39,
43100 Parma, Italy
e-mail: giacomo.rizzolatti@unipr.it
M. Fabbri-Destro
e-mail: fbbmdl@unife.it
G. Rizzolatti
Istituto Italiano di Tecnologia (IIT) Unità di Parma,
Parma, Italy
M. Fabbri-Destro
Dipartimento SBTA, Sezione di Fisiologia Umana,
Università di Ferrara, via Fossato di Mortara,
17-19, 44100 Ferrara, Italy
Our article appeared in Experimental Brain Research a few
months later.
The idea of sending our report on mirror neurons to
Experimental Brain Research, rather than to another neuro-
science journal, was motivated by a previous positive expe-
rience with that journal. A few years earlier, Experimental
Brain Research accepted an article in which we presented
(Rizzolatti et al. 1988) a new view (something that typi-
cally referees did not like) on the organization of the ventral
premotor cortex of the monkey and reported the Wndings
that paved the way for the discovery of mirror neurons. In
that article we described how, in the ventral premotor cor-
tex (area F5) of the monkey, there are neurons that respond
both when the monkey performs a motor act (e.g., grasping
or holding) and when it observes an object whose physical
features Wt the type of grip coded by that neuron (e.g., pre-
cision grip/small objects; whole hand/large objects). These
neurons (now known as “canonical neurons”, Murata et al.
1997) and neurons with similar properties, described by
Sakata et al. (1995) in the parietal cortex are now univer-
sally considered the neural substrate of the mechanism
through which object aVordances are translated into motor
acts (see Jeannerod et al. 1995).
We performed the experiments on the motor properties
of F5 in 1988 using an approach that should almost neces-
sarily lead to the discovery of mirror neurons if these neu-
rons existed in area F5. In order to test the F5 neurons with
objects that may interest the monkeys, we used pieces of
food of diVerent size and shape. To give the monkey some
food, we had, of course, to grasp it. To our surprise we
found that some F5 neurons discharged not when the mon-
key looked at the food, but when the experimenter grasped
it. The mirror mechanism was discovered.
The next important role of Experimental Brain Research
in the discovery of mirror neurons was its acceptance in
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Fig. 1 Example of an F5 mirror neuron selectively discharging during
monkey grasping movements and during observation of a grasping
movement done by the experimenter. a Lateral view of the brain with
indicated the location of F5. b Grasping observation. c Grasping exe-
cution. a arcuate sulcus, c central sulcus, ip intraparietal sulcus (from
di Pellegrino et al. 1992)
1996 of two articles which Wrst reported the existence of the
mirror areas in humans (Rizzolatti et al. 1996; Grafton et al.
1996). The rational of the experiment was as follows: If
mirror mechanism exists in humans the observation of
actions done by another individual should activate, besides
visual areas, also areas that have motor properties. We ran
two PET experiments and showed that indeed the areas
where the mirror neurons are located in the monkey
become also active in humans. This Wnding was subse-
quently replicated by dozens of experiments (see Rizzolatti
and Craighero 2004; Rizzolatti et al. 2009; Cattaneo and
Rizzolatti 2009).
At present there is an enormous literature on mirror
neurons. A set of it concerns experiments in monkeys (see
Rizzolatti and Craighero 2004; Rizzolatti et al. 2009) and
more recently in birds (Prather et al. 2008); another set,
much larger, concerns experiments in humans. In the pres-
ent article we (GR and MF-D) will review mirror data in
humans, examining, however, (by necessity) only part of
the enormous mirror neuron studies triggered by our initial
PET studies published in Experimental Brain Research.
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Exp Brain Res (2010) 200:223–237
The organization of the human parieto-frontal mirror
system
In humans the observation of goal-directed motor-acts acti-
vates, besides visual areas, the inferior parietal lobule (IPL)
and the premotor cortex, mostly its ventral part, plus the
caudal part of the inferior frontal gyrus (IFG) roughly cor-
responding to the pars opercularis of Broca’s area. These
two regions form the core of the human parieto-frontal mir-
ror system (Rizzolatti and Craighero 2004; Fabbri-Destro
and Rizzolatti 2008) (Fig. 2).
Both the premotor and parietal nodes of the human mir-
ror system present a somatotopic organization, albeit rather
rough (Buccino et al. 2001; Wheaton et al. 2004; Sakreida
et al. 2005; Etzel et al. 2008). Observation of motor acts
done by others with the leg, hand, and mouth activates the
precentral gyrus and the pars opercularis of IFG in a
medial to lateral direction, as in the classical homunculus of
PenWeld and Rasmussen (1950)and Woolsey et al. (1952).
In IPL, mouth motor acts appear to be represented rostrally,
hand/arm motor acts caudally and leg motor acts even more
caudally, and dorsally extending into the superior parietal
lobule. A similar somatotopic organization based on the
motor properties of the recorded neurons has been recently
reported in monkey IPL by Rozzi et al. (2008) (see also
Hyvarinen 1982).
It is open question whether the activations found during
the observation of reaching to grasp movements around the
superior frontal sulcus (e.g., Grèzes et al. 2003; Buccino
et al. 2004a; Gazzola and Keysers 2009) are due to repre-
sentation of proximal movements or to motor preparation.
This uncertainty depends of the fact that there is no clear
boundary between the ventral (PMv) and dorsal (PMd) pre-
motor cortices in humans. According to the Wrst, somato-
topic, interpretation, the dorsal premotor activation is
Fig. 2 Later view of human brain. The colored areas form the pari-
eto-frontal mirror network. Red parietal mirror node, yellow frontal
mirror node
Exp Brain Res (2010) 200:223–237
located in a speciWc sector of PMv where proximal move-
ments are represented. According to the motor preparation
interpretation, the dorsal premotor activations are located in
PMd, an area that according to monkey single neuron data
is mostly involved in covert motor preparation (Kalaska
and Crammond 1995; Crammond and Kalaska 2000).
The parietal region active during the observation of
object-directed motor acts is mostly located in the sector of
the IPL close to and inside the intraparietal sulcus. This
restricted localization raises an interesting question: Are
other types of motor act also represented in other parts of
IPL and, in that case, what kinds of act? An answer to these
questions has been recently obtained by two fMRI studies.
The Wrst investigated the localization of intransitive move-
ments (Lui et al. 2008), the other that of actions performed
with tools (Peeters et al. 2009).
In the Wrst study, volunteers were scanned while they
observed mimed, symbolic, and meaningless motor acts. As
during the observation of object-directed actions, fMRI sig-
nal increase was found in the premotor cortex and in IPL.
However, while the premotor cortex activation overlapped
that previously found during the observation of object-
directed actions, in the parietal lobe the signal increase was
not restricted to the intraparietal sulcus region, but extended
into the posterior part of the supramarginal gyrus and the
angular gyrus. Most interestingly, while the mimed actions
were located dorsally close to the intraparietal sulcus, that
is in a location similar to that activated by the observation
of actual object-directed movements, symbolic motor acts
were located ventrally, mostly in the angular gyrus. (Lui
et al. 2008).
In the second study, volunteers observed a variety of
motor acts performed by another individual either by hand
or using tools. The results showed that the observation of
motor acts performed with tools activates the parieto-fron-
tal circuit mediating hand grasping and, in addition, a spe-
ciWc sector of the anterior part of the left supramarginal
gyrus (aSMG). In a parallel experiment carried out in the
same study on naïve as well as on monkeys proWcient in
using tools (rake and pliers), no evidence was found for a
parietal sector activated during tool action observation. It
was concluded that aSMG is a new evolutionary acquisition
of homo sapiens that mediates the human capacity to under-
stand the causal relationship between tools and the goal of
the action achieved by using tools.
Typically, an activation of SPL is absent or marginal in
those studies where the experimenters use as visual stimuli
distal motor acts or acts in which the distal component is
prominent. The possibility of a proximal mirror represen-
tation in SPL was recently tested in an fMRI study where
volunteers were asked to transport their hand to a partic-
ular location in space without grasping objects. The
reaching movements were executed, observed, or imagined.
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An overlap between executed, observed, and imagined
reaching activation was found in SPL extending into IPS,
and in PMd. This study provides the Wrst demonstration of
a mirror mechanism for reaching movements (Filimon et al.
2007).
Evidence for the activation of human cortical motor
system during the observation of actions done by others
The brain imaging studies reviewed above show that human
cortical areas, active when individuals watch actions done
by others, strictly correspond to the cortical areas that are
endowed with mirror properties in the monkey (Rizzolatti
et al. 2009; for monkey fMRI data see Nelissen et al. 2005).
Because mirror neurons are motor neurons, the observation
of motor acts done by others should determine, if mirror
neurons are present in humans, an increase of motor cortex
excitability congruent with the observed motor act.
Evidence that this is the case has been obtained using
transcranial magnetic stimulation (TMS). Fadiga et al.
(1995) recorded the motor-evoked potentials (MEPs)
induced by the stimulation of the left motor cortex in vari-
ous muscles of the right hands and arms of volunteers asked
to watch an experimenter while he grasped objects with his
hand or performed meaningless arm movements. As a con-
trol for attentional factors there was a third condition in
which volunteers detected the dimming of a small light.
During both the experimental conditions there was a clear
increase in the observer’s MEPs, relative to the control con-
dition. This increase was present in those muscles that were
recruited when the tested individuals were asked to execute
the observed movements. Several TMS experiments con-
Wrmed these Wndings (e.g., Strafella and Paus 2000; Gangitano
et al. 2001; Maeda et al. 2002; Borroni et al. 2005). Among
them particularly interesting is the study by Gangitano et al.
(2001). These authors showed not only that MEPs recorded
from the hand muscles increased during grasping observa-
tion, but also that the relative cortical facilitation closely
reXected the diVerent grasping phases (Fig. 3).
EEG and MEG studies provided further evidence of acti-
vation of the motor cortex during action observation.
Already, in the 1950s, Gastaut and Bert (1954) showed that
the
rhythm, a rhythm recorded in the correspondence of
the cortical motor areas and known to desynchronize during
movement execution, also desynchronizes during the obser-
vation of actions carried out by others. Following the dis-
covery of mirror neurons, several studies (e.g., Altschuler
et al. 1997; Cochin et al. 1999) repeated these experiments
conWrming the desynchronization of
rhythm during
action observation.
Similar results were also obtained using magneto-
encephalography (MEG) (Hari et al. 1998), a technique that
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Fig. 3 Modulation of the motor cortex excitability during grasping
observation. a Schematic sequence of events during a grasping
trial. b Averaged values of motor-evoked potentials (MEPs) of a hand
muscle (Wrst dorsal interosseus) collected at diVerent times during the
analyses the brain electric activity on the basis of the mag-
netic Welds it generates. MEG data provided evidence of a
desynchronization of the cortical rhythms of observer’s
motor cortex (including those originating from the cortex
located inside the central sulcus) during object manipula-
tion and when the manipulation was observed.
Mirror activity is modulated by motor experience
There is evidence that only motor acts that are present in
the motor repertoire of the observer are eVective in activat-
ing the mirror neuron system. In an fMRI experiment nor-
mal volunteers observed video-clips showing mouth motor
acts made by humans, monkeys, and dogs. In one condi-
tion, the observed motor act was biting, a motor act present
in the motor repertoire of all three species and in another
condition the stimuli were communicative gestures proper
to each species: reading silently a text, lip-smacking, and
barking. The data demonstrated that the left IPL and IFG
responded to actions made by human and nonhuman per-
formers, as long as the action was part of the human motor
repertoire (e.g., biting). In contrast, there was no activation
(barking) or almost no activation (lip-smacking) when the
action belonged to another species (Buccino et al. 2004b).
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Exp Brain Res (2010) 200:223–237
observation of grasping movements. 500 ms: hand at the starting
position (time value refers to the onset of the video clip showing the
action), 3,000 ms: hand maximum aperture (from Gangitano et al.
2001)
Mirror neuron activation is also related to the observer’s
motor experience of a given action. This has been nicely
demonstrated in experiments using dance steps as observed
stimuli. First it was shown that, in the observer, the amount
of mirror activation correlated with the degree of the
observer’s motor skill for that action (Calvo-Merino et al.
2005). A further experiment ruled out the possibility that
this eVect could be due to mere visual familiarity with the
stimuli. The observation of steps that are peculiar to male
dancers determined a stronger mirror activation in male
professional dancers than those performed by female danc-
ers and vice versa (Calvo-Merino et al. 2006). A further
prospective study showed that dancers initially naïve to cer-
tain steps showed an increase in mirror activation over time
if they underwent a period of motor training in which they
became skillful in performing the same steps (Cross et al.
2006).
Some clues to the mechanism responsible for these
eVects come from experiments that tested whether conver-
gence of observation and execution of motor acts facilitates
the building of motor memories. These experiments
showed that after a training period in which participants
simultaneously performed and observed congruent move-
ments there was a potentiation of the learning eVect, with
respect to motor training alone, as shown by the kinematics
Exp Brain Res (2010) 200:223–237
of the movement evoked by TMS (Stefan et al. 2005,
2008). Further evidence in favor of plasticity of the mirror
mechanism comes form experiments showing that the mir-
ror responses triggered by a corresponding movement
could be modiWed by repetitively coupling the performed
movement with the observation of diVerent movements
(Catmur et al. 2007, 2008).
The functional roles of mirror neurons
Mirror neurons, directly recorded or demonstrated by non-
invasive techniques, are present in various cortical areas of
primates (Rizzolatti and Craighero 2004) and in birds
(Prather et al. 2008). All of them are endowed with the
same mechanism: a mechanism that translates sensory
information describing motor acts done by others into a
motor format similar to that the observers themselves gen-
erate when they perform those acts.
While the mirror mechanism is the same regardless of
the location of neurons endowed with it, the result of the
sensory-motor transformation depends on the location of
mirror neurons. Those located in emotional centers like the
insula or the cingulate cortex intervene in phenomena like
empathy (see Gallese et al. 2004), while those located in
the parieto-frontal circuit provides the observer with motor
representations of others’ motor actions devoid of emo-
tional content (Rizzolatti and Craighero 2004).
From the discovery of mirror neurons in area F5 of the
monkey, two explanations, not mutually exclusive, have
been proposed for the functional role of the mirror neurons
in this area and in the IPL. The Wrst was that mirror neurons
underlie imitation. The second was that the correspondence
between the motor format generated by observing others
and that generated internally in order to act enables the
observer to understand others’ behavior, without the neces-
sity for complex cognitive elaborations.
The Wrst view has been thought of as unlikely because of
ethological data showing that monkeys, unlike humans and
apes, do not imitate (Visalberghi and Fragaszy 1990). As a
matter of fact “imitation” phenomena are also present in
“lower” primates (see Zentall 2006). For example, tongue
protrusion in response to the same motor act done by
another individual, described many years ago by MeltzoV
and Moore (1979) in newborn babies, has been recently
reported in macaque monkeys (Ferrari et al. 2006). Yet,
convincing evidence of “true imitation”, that is imitation in
which the learned behavior is performed with the same
movements (including hand movements) as shown by the
teacher is lacking. Thus, the presence of a well-developed
mirror mechanism concerning hand movements suggests that
understanding motor acts done by others, rather than imita-
tion, is the evolutionary older function of the parieto-frontal
227
mirror network in the monkey. Other cognitive functions
like imitation (see below) and, most likely, language
(Rizzolatti and Arbib 1998) evolved on the top of it.
In the present review we will discuss Wrst the relations
between mirror mechanism and imitation. We will deal
later with goal and intention understanding, because the
discussion of neurophysiological mechanisms underlying
these capacities leads directly to the issue of autism, whose
discussion will conclude this article.
Mirror mechanism and Imitation
The term imitation has many deWnitions in human litera-
ture. There are, however, two main senses in which it is
most commonly used (see Rizzolatti 2005). The Wrst deW-
nes imitation as the capacity of an individual to replicate an
observed motor act; the second deWnes imitation as the
capacity to acquire, by observation, a new motor behavior
and to repeat it using the same movements employed by the
teacher. In both cases imitation requires the capacity to
transform sensory information into a motor representation
of it.
There is convincing evidence that the mirror mechanism
is involved in imitation as an immediate replica of the
observed motor act. In an fMRI experiment, volunteers
were tested in two main conditions: “observation” and
“observation–execution”. In the “observation” condition,
participants were shown a moving Wnger, a cross on a sta-
tionary Wnger, or a cross on empty background. The
instruction was to observe the stimuli. In the “observation–
execution” condition, the same stimuli were presented, but,
this time, the instruction was to lift the right Wnger, as fast
as possible, in response to them. The crucial contrast was
between the trials in which the volunteers made the move-
ment in response to an observed action (“imitation”) and
the trials in which the movement was triggered by the cross
(a non-imitative behavior). The results showed that the acti-
vation of the mirror system and in particular of the posterior
part of IFG was stronger during “imitation” than in other
conditions (Iacoboni et al. 1999).
Further evidence that the mirror system plays a crucial
role in this kind of imitation was provided by repetitive
TMS (rTMS), a technique that determines a transient
depression of the stimulated region. In a group of volun-
teers the caudal part of the left frontal gyrus (Broca’s area)
was stimulated while they (a) pressed keys on a keyboard,
(b) pressed the keys in response to a point of red light indi-
cating which key to press, (c) imitated a key pressing
movement done by another individual. The data showed
that rTMS lowered the participants’ performance during
imitation, but not during the other two tasks (Heiser et al.
2003).
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228 Exp Brain Res (2010) 200:223–237

Evidence that mirror system is involved in imitation
learning comes both from EEG and fMRI studies. Marshall
et al. (2009) examined diVerences in EEG desynchroniza-
tion during observation of drawing of various characters
selected from the Cham alphabet, an alphabet used in
Southeast Asia, and with which none of the participants
was familiar. Compared to carrying out unrelated drawing
(Latin letters), brief imitative experience was speciWcally
associated with a signiWcantly larger desynchronization in
the 11–13 Hz band at mid-frontal sites (F3 and F4) when a
previously imitated action was presented again. In addition,
higher Wdelity of imitation was signiWcantly correlated with
greater bilateral desynchronization of the
rhythm at cen-
tral sites (C3 and C4) during subsequent observation of the
previously imitated action.
A more elaborate experimental design was used by
Buccino et al. (2004a). Using an event-related fMRI
paradigm these authors tested musically naive participants
during four events: (1) observation of guitar chords played
by a guitarist, (2) a pause following model observation, (3)
execution of the observed chords, and (4) rest. The results
showed that the basic circuit underlying imitation learning
consists of the IPL and the posterior part of IFG plus the
adjacent premotor cortex. This circuit starts to be active
during the Wrst event: observation. During pause, i.e.,
during the phase in which visual information is elaborated
for action production, activations are observed in the
middle frontal gyrus (area 46) and in structures involved in
motor preparation (dorsal premotor cortex, superior parietal
lobule, rostral mesial areas). The activation of these areas
plus the somatosensory and motor areas contralateral to the
hand used to execute chords dominates the subsequent
execution phase (Fig. 4).
On the basis of this experiment and a following one also
based on learning of playing guitar chords (Vogt et al.
2007), the authors proposed a model of imitation learning
(see Byrne 2002 for a similar model-based ethological
observations) consisting of two distinct processes: (a)

Fig. 4 Cortical activations dur-

ing imitation learning. Upper
part graphic illustration of the
events forming the experimental
conditions imitation (IMI) and
non-imitation (NON IMI). Both
conditions consisted of four
events preceded by the presenta-
tion of a colored cue (a square)
informing the participants on the
task they have to perform. IMI
condition: event 1 observe the
teacher’s hand playing the chord
(IMI-1), event 2 rehearse the
observed chord (IMI-2), event 3
replicate it. Event 4 keep the
hand still. NON IMI condition:
event 1 observe the teacher’s
hand playing the chord (Non
IMI-1), event 2 do not rehearse
the observed chord (Non IMI-2),
event 3 touch the neck of the
guitar, without playing a chord
(Non IMI-3). Event 4 keep the
hand still. Lower part cortical
areas activated during events 1
and 2 in IMI and Non IMI
conditions (modiWed from
Buccino et al. 2004a, b)

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Exp Brain Res (2010) 200:223–237
segmentation of the action to be imitated into its individual
elements and their transformation into the corresponding
potential movements of the observer; (b) organization of
these potential movements into a temporal and spatial pat-
tern that replicates that shown by the demonstrator. The
Wrst process is achieved through the mirror mechanism,
while the second one is mostly due to the activity of the
prefrontal lobe and in particular of area 46 that memorizes
and recombines the motor elements in the new pattern.
Action and intention understanding
Coding the goals of the motor acts
Social life is based on our capacity to understand the behav-
ior of others. Let us imagine this situation. John and Mary
are in a pub and John’s hand comes into contact with a mug
of beer; Mary immediately understands whether he is
grasping it or not. Moreover, according to how he grasped
it, she can also understand why he is doing it (e.g., for
drinking or for giving the mug to a friend). How does Mary
understand the goal of the John’s motor act and the inten-
tion behind it?
One possibility is that she is using an inferential reason-
ing elaborating the acquired visual information through
some cognitive mechanism (see Frith and Frith 1999;
Csibra and Gergely 2007). Another possibility is that this is
not necessary in this simple situation, and the understand-
ing of what John is doing and why he is doing it, is acquired
through a mechanism that directly transforms visual
information into a motor format. The proprieties of mirror
neurons support the existence of such a mechanism.
There is clear evidence from monkey experiments that
neurons in the parietal lobe, premotor cortex, and even in
the primary motor cortex, code the goal of a motor act
rather than, as traditionally thought, movements of body
parts (Rizzolatti et al. 1988; Kakei et al. 1999, 2001;
Fogassi et al. 2005; Umiltà et al. 2008). Mirror neurons
located in F5 and in IPL have motor properties identical to
those of purely motor neurons. Thus, because the electrical
activity recorded in these experiments during voluntary
behavior and action observation always consists of action
potentials (the neuron output) the messages conveyed dur-
ing voluntary movement and during mirror activation are
identical. In both cases the neurons send information on the
goal of the observed motor act.
Given these Wndings, it appears logical to assume that a
similar organization does exist also in humans. Evidence in
this sense came from fMRI studies. Gazzola et al. (2007a)
instructed volunteers to observe video-clips where either a
human or a robot arm grasped objects. In spite of diVer-
ences in shape and kinematics between the human and
229
robot arms, the parieto-frontal mirror system was activated
in both conditions. These data was recently conWrmed and
extended by Peeters et al. (2009).
Further evidence in favor of goal coding in the human
mirror network comes from an fMRI study in which indi-
viduals were tested both during motor execution and when
listening to the sound of an action made by the same eVec-
tor (Gazzola et al. 2006). The results showed, in both cases,
a similar activation of the left parieto-frontal circuit.
An experiment on aplasic individuals conWrmed that the
mirror network codes the goal of motor acts (Gazzola et al.
2007b. In this study the authors addressed the following
question: Can the goal of a hand movement be recognized
in the absence of any experience of hand movements? To
answer it two individuals born without arms and hands
were studied. While being scanned they were asked to
watch video-clips showing hand actions and their brain
activations were compared with those of control volunteers.
All participants also made actions with diVerent eVectors
(feet, mouth and, for normal volunteers, hands). The results
showed that the mirror system of aplasic individuals was
activated by the observation of hand motor acts. This dem-
onstrates that the brains of aplasics can mirror motor acts
that they have never executed. The goal is recognized
through the recruitment of areas involved in the execution
of motor acts having the same goal but using diVerent eVec-
tors.
The issue of goal coding was recently addressed by
Hamilton and Grafton (2006) using the adaptation tech-
nique, a technique based on the trial-by-trial reduction of a
physiological response to repeated stimuli. Participants
observed a series of video-clips showing goal-directed
motor acts with the sequence controlled so that some goals
were novel and others repeated relative to the previous
movements. Repeated presentation of the same goal caused
the suppression of the response in the left intraparietal sul-
cus (IPS) while this region was not sensitive to the trajec-
tory of the actor’s hand.
While the fMRI data support, in agreement with monkey
data, the notion that mirror neurons code motor acts, most
TMS data appear to indicate that during the observation of
motor acts performed by others, there is an activation of the
neural substrate controlling the muscles that are involved in
that motor act (see Rizzolatti and Craighero 2004). There is
an ingenious study, however, that was able to demonstrate,
using TMS, goal coding in human motor cortex (Gangitano
et al. 2001). In this study motor-cortex excitability was
tested during the observation of hand movements directed
to a speciWc goal (predictable movements) and in trials in
which the hand moved in a diVerent direction (unpredict-
able movements). The data showed that the observation of
unpredictable movements did not elicit the expected change
in the excitability of the motor cortex corresponding to the
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observed movements. During the observation of the unpre-
dictable movements, the excitability pattern was the same
as that found during the observation of the predicted ones.
This indicates that the observed motor acts were coded,
from their very beginning, in terms of the Wnal goal of the
action and not in terms of the movements forming them.
Coding the intention behind the motor acts of others
Studies of the mirror mechanism have provided evidence
for its role in motor act understanding. What about inten-
tion understanding? Evidence that the mirror mechanism
play a role in this capacity has been Wrst provided by an
fMRI study. In this study there were three conditions. In the
Wrst one (“context”) the volunteers saw some objects (a tea-
pot, a mug, a plate with some food on it) arranged as if a
person was ready to drink the tea or as if a person had just
Wnished having his/her breakfast; in the second condition
(“action”) the volunteers were shown a hand that grasped a
mug without any context; in the third condition (“inten-
tion”) the volunteers saw the same hand action within the
previous two contexts. The context and the diVerent grip
shapes suggested the intention of the agent, i.e., grasping
the cup for drinking or grasping it for cleaning the table
(Iacoboni et al. 2005).
The results showed that in both action and intention con-
ditions there was an activation of the mirror mechanism.
Crucial was the comparison between intention and action
conditions. This comparison showed that the understanding
of the intention of the doer determined a marked increase in
the activity of the mirror mechanism.
The importance of the mirror system in understanding
intention has been recently conWrmed by an fMRI experi-
ment based on the adaptation paradigm. Participants were
asked to observe repeated movies showing either the same
movement or the same outcome independent of the exe-
cuted movement. The results showed activity suppression
in the right inferior parietal and in the right IFG when the
outcome was the same. Kinematic parameters do not appear
to inXuence the activity of these regions. These Wnding
indicate therefore that the right hemisphere mirror system
encodes the physical outcomes of human actions, an initial
step for inferring intentions underlying these actions
(Hamilton and Grafton 2008).
Taken together, these data suggest that the intentions
behind the actions of others can be recognized by the mirror
neuron mechanism. These Wndings do not imply that other
more cognitive ways of “reading minds” do not exist.
Indeed, recent fMRI studies showed that, in speciWc condi-
tions, the understanding of motor acts performed by others
might require, beside the mirror mechanism, the activation
of areas outside those forming the mirror system. For
example, when tasks require a top–down inference either to
123
Exp Brain Res (2010) 200:223–237
assess the meaning of a motor act in an implausible context
(Brass et al. 2007; Liepelt et al. 2008) or to judge whether
the intention of the observed action was ordinary or unusual
(de Lange et al. 2008), there is an increase of activity in the
posterior superior temporal sulcus (STS) region, posterior
cingulate cortex, and the medial prefrontal cortex.
Some studies suggested that a region of right temporo-
parietal junction, often referred to as right TPJ, plays a cru-
cial role in ‘mentalizing’ (e.g., Saxe and Wexler 2005,
2006). This view, however, should be accepted with cau-
tion. In fact, as shown by Mitchell (2008) (see, however,
Scholz et al. 2009) the same right TPJ activated during
“mentalizing” is also active in task requiring attention. The
overlap between these two mental functions casts serious
doubts on the hypothesis that TPJ plays a crucial role in
intention understanding.
In accord with the interpretation of Mitchell’s view are
some data by Buccino et al. (2007). In an fMRI study these
authors investigated the neural basis of human capacity to
diVerentiate between actions reXecting the intention of the
agent (intended actions) and actions that did not reXect it
(non-intended actions). Volunteers were presented with
video-clips showing a large number of actions performed
with diVerent eVectors, each in a double version: one in
which the actor achieved the purpose of his or her action
(e.g., pour the wine), the other in which the actor performed
a similar action but failed to reach the goal of it because of
a motor slip or a clumsy movement (e.g., spill the wine).
The data showed the activation of the mirror system areas
in both conditions. The contrast, however, non-intended
versus intended actions showed an activation of the right
TPJ and the mesial prefrontal cortex. Because there is little
doubt that a person observing another person falling down
or spilling the wine because of a motor slip does not “put
himself in the shoes” of that individual, the activation
observed in the experiment by Buccino et al. (2007) is
hardly due to an attempt to understand the other’s intention,
but rather depends on an increase of the observer’s atten-
tion due to the surprising course of the event.
An impairment of the mirror mechanism explains some
deWcits in children with autism
Autistic spectrum disorder (ASD) is a heterogeneous syn-
drome characterized by impairment in social skills, verbal
and nonverbal communication, coupled with restricted, and
repetitive behaviors (DSM-IV-TR 2000). DeWcits in the
domains of aVective links and emotional behavior are other
aspects of ASD (Kanner 1943).
Autism aVects a variety of nervous structures, from the
cerebral cortex to the cerebellum and brainstem (see
Minshew and Williams 2007). However, in the context of a
Exp Brain Res (2010) 200:223–237
broader neurodevelopmental deWcit, a set of ASD symp-
toms (impairment in communication, language, and the
capacity to understand others) appears to match functions
mediated by the mirror mechanism. The hypothesis that
this speciWc set of deWcits might depend on an impairment
of the mirror mechanism (Altschuler et al. 2000; Williams
et al. 2001) has therefore been advanced.
Evidence coming from EEG, TMS, and brain imaging
studies supports this hypothesis (e.g., Nishitani et al. 2004;
Oberman et al. 2005; Theoret et al. 2005; Dapretto et al.
2006). Oberman et al. (2005) studied the suppression of

rhythm during the execution and observation of motor acts
in typically developing (TD) and children with autism. The
results showed that, in contrast with TD children, ASD do
not present
rhythm suppression during the observation of
motor acts done by others. The
rhythm suppression is
present only during active movements (Fig. 5). Similar data
were obtained by Martineau et al. (2008).
Additional evidence for an impaired mirror mechanism
in autism came from behavioral and TMS studies. Avikainen
et al. (2003) showed that, unlike TD individuals, who,
when viewing persons face-to-face, tend to imitate them in
a mirror way, children with autism do not show this prefer-
ence. This imitation peculiarity is most likely due to a deW-
cit of mirror mechanism coding other person’s movements
on one’s own. Theoret et al. (2005) demonstrated an
impaired motor facilitation in children with autism during
action observation by using TMS.
Finally, strong evidence in favor of a deWcit of the mirror
mechanism in ASD came from an fMRI study. High
functioning children with autism and matched controls
were scanned while imitating and observing emotional
Fig. 5 Absence of EEG desynchronization during the observation of
movements done by others. The charts show desynchronization of the

rhythm in controls (a) and patients with autism spectrum disorder
(b). Observation of movement of an inanimate object (pale green),
movements made with the hand (green), and active hand movements
(red). The bars represent the amount of
activity in central scalp
231
expressions. The results showed a signiWcantly weaker acti-
vation in IFG in children with autism than in typically
developing (TD) children. Most interestingly, the activation
was inversely related to symptom severity (Dapretto et al.
2006).
Taken together, these data indicate that children with
autism process the actions done by others in a manner
diVerent from that of TD children. The simplest way to
account for these diVerences is to postulate (see also above)
that children with ASD have an impairment of the mirror
mechanism. This hypothesis is also known as the “broken
mirror” hypothesis (Ramachandran and Oberman 2006).
There are some behavioral studies indicating, however,
that this hypothesis is not fully satisfactory and needs spec-
iWcations. These studies reported that children with ASD do
not present deWcits in understanding the goal of motor acts
done by others (Hamilton et al. 2007; Bird et al. 2007;
Leighton et al. 2008; Southgate and Hamilton 2008). It
was, therefore, claimed that the “broken mirror” hypothesis
of autism is wrong (e.g., Southgate and Hamilton 2008).
It must be noted, however, that these studies took into
account only one aspect of mirror organization, the one
related to the role of the mirror neurons in the recognition
of motor acts done by others. If only this aspect of the mir-
ror system is considered, the criticism against the broken
mirror hypothesis appears to be well taken.
Neurophysiological studies showed, however, that there
is a second aspect of the mirror neuron organization based
not on the activity of single neurons, but on the organiza-
tion of cortical motor system. The neural basis of this orga-
nization consists of chains of motor acts. These chains are
formed by populations of neurons, each coding speciWc,
locations; C3, Cz, and C4 refer to scalp coordinates of the 10/20 EEG
system. SigniWcant
activity, indicated by asterisks, is present for the
hand observation condition only in controls, showing that patients with
autism spectrum disorder fail to react to the observation of other
people’s actions in the standard way (modiWed from Oberman et al.
2005)
123
232 Exp Brain Res (2010) 200:223–237

serially connected motor acts (e.g., reach-grasp-bring to the
mouth; or reach-grasp-move away). During voluntary
movements the agent recruits one of these chains according
to his/her motor intention. These chains also contain
“action constrained” mirror neurons, that is neurons that
Wre only if the motor act they code is part of a motor chain
(e.g., grasping for placing, but not grasping for eating, or
viceversa). During the observation of actions done by oth-
ers, “action constrained” mirror neurons Wre when the
observed behavior matches the speciWc action coded by the
chain in which those neurons are embedded. Their Wring
activates an entire action chain providing the observer with
a motor representation of the action that the agent is ready
to do. In virtue of this mechanism the observer understands
the agents’ intention (Fogassi et al. 2005).
Recently, it has been shown that the chained motor act
organization is impaired in autism (Cattaneo et al. 2007).
TD children and children with autism were asked to per-
form the two actions: grasping an object to eat it or grasp-
ing to place it into a container (Fig. 6). The EMG activity of
the mylohyoid muscle (MH), a muscle involved in opening
of the mouth, was recorded. In TD children the muscle
became active as soon they moved the arm to reach the
food. In contrast, no MH muscle activation was observed
during food reaching and grasping in autistic children. MH
muscle activation appeared only when the children brought
the food to their mouth. These data indicate that ASD chil-
dren are unable to organize their motor acts into a unitary
action characterized by a speciWc intention.
In a further experiment TD children and children with
autism were tested while they observed an experimenter
either grasping a piece of food for eating or grasping a
piece of paper for placing it into a container (Fig. 6). The
EMG of MH muscle was recorded. The results showed that
in TD children, the observation of food grasping
determined the activation of MH, while this activation was
lacking in children with autism. In other words, while the
observation of an action done by another individual
intruded into the motor system of a TD observer, this
intrusion was lacking in children with autism. This Wnding

Fig. 6 Responses of typically

developing children (TD) and
children with autism (AU) dur-
ing the observation and execu-
tion of two actions. Upper panel
schematic representation of the
actions. Top the individual
reaches a piece of food located
on a touch-sensitive plate, grasps
it and brings it to the mouth.
Bottom the individual reaches a
piece of a paper located on the
same plate, grasps it, and puts
into a container placed on the
shoulder. Middle panel time
course of the rectiWed EMG
activity of MH muscle for TD
children and children with
autism (AU) during the
observation of the two tasks.
Lower panel EMG activity of
MH muscle during the execution
of the two tasks. Bringing-to-
the-mouth action (red), placing
action (blue). Vertical bars
indicate the SE. All curves are
aligned with the moment of
object lifting from the touch-
sensitive plate (t = 0, dashed
vertical line) (modiWed from
Cattaneo et al. 2007)

123
Exp Brain Res (2010) 200:223–237
indicates that, in autism, the mirror system is silent during
action observation and the immediate, experiential under-
standing of others’ intention is absent.
Summing up, these data strongly suggest that children
with autism have a deWcit in the chained organization of
motor acts and, as a consequence, they are unable to acti-
vate it during action observation. Without this internal
“replica” of the actions of others, they cannot grasp
directly, without cognitive inferences, the intention of oth-
ers.
According to this new mirror hypothesis on the neural
basis of the autistic deWcit in understanding others, there is
a dissociation, in autism, between the capacity to under-
stand the what of an action (carried out by the basic mirror
neurons mechanism) and the why of it (depending on the
integrity of the chained motor organization). In order to test
this point an experiment was recently performed by Boria
et al. (2009). In this study TD and autistic children were
presented with pictures showing goal-directed motor acts
and asked to report what the actor was doing and why he
was doing it. These two tasks test two diVerent abilities: the
Wrst is that of recognizing the goal of the observed motor
act, that is something occurring “now” (e.g., grasping an
object); the second consists in understanding the intention
of the action that is something that will occur in the future
(e.g., grasping to eat). The results showed that while both
TD and ASD children recognized what the actor was doing,
ASD children were impaired in recognizing the why of that
action. The type of why error was systematic: ASD chil-
dren constantly tended to attribute to the actor the intention
related to the common use of the grasped object. Thus,
grasping a pair of scissors indicates the intention to cut,
while grasping a mug that of drinking, this regardless of
how the object was grasped. In other words, ASD children
interpreted the behavior of others on the basis of the com-
mon use of observed object rather then on the basis of the
motor behavior of the actors (Boria et al. 2009).
How the things are now
I (GR) suppose that none of the authors of the Wrst short
report on mirror neurons (Di Pellegrino et al. 1992) would
have predicted the enormous impact that that discovery
would have not only on neuroscience but also on a host of
disciplines ranging from social sciences to esthetics. What
is particularly rewarding for who discovered mirror neu-
rons is that, after so many years, new important Wndings
related to the mirror mechanism continue to appear. Three
recent particularly interesting contributions on mirror
mechanisms in the monkey are worth mentioning.
The Wrst shows that, besides mirror neurons describing
the motor acts done by others independently of their
233
distance from the observer, there are other mirror neurons
in area F5 that discharge depending on whether the motor
act is performed within or outside the monkey’s periper-
sonal space. Most interestingly, some of these neurons code
space operationally. That is, these neurons describe the
action of others in term of their possibility to act on an
object. For a set of these neurons, for example, the periper-
sonal space becomes extrapersonal if a transparent barrier
is placed between the observing monkey and the stimulus
(Caggiano et al. 2009).
A second recent study showed that a subset of neurons,
located in area LIP, “mirrors” observed attention by Wring
both when monkey looks in the preferred direction of the
neuron and when the observed monkey looks in that direc-
tion. Another subset of LIP of neurons was, in contrast,
suppressed by social gaze cues, possibly subserving behav-
ioral demands by maintaining Wxation on the observed face.
As proposed by the authors (Shepherd et al. 2009), these
Wndings suggest that the mirror mechanism of area LIP
contributes to sharing of observed attention, a fundamental
step in social cognition.
Finally a third study showed that, in the monkey ventral
intraparietal area (VIP) and the adjacent area PFG there are
neurons that code the peripersonal space of the observing
(recorded) monkey and the peripersonal space of another
individual facing it (Ishida et al. 2009). This Wnding sug-
gests that mirror neurons are critical for understanding not
only the motor acts of others, but also others’ body parts
and body-centered motor acts.
As one can imagine, a new view of the neural basis of
cognition has also raised doubts and criticisms. Some
of them have been very useful for clarifying various points
of the “immediate action perception” theory and, more gen-
erally, for sharpening and rendering more precise its claims
(e.g., Jacob and Jeannerod 2005; Knoblich and Prinz 2005;
Csibra and Gergely 2007).
Beside these constructive criticisms, recently some arti-
cles appeared expressing a curious “anti-mirror” stance
(Dinstein 2008; Dinstein et al. 2008; Lingnau et al. 2009).
Their main argument is the following: action observation
activates in humans, as in the monkeys, parietal and motor
areas, but the properties of the activated neurons are diVer-
ent in the two species. While in monkeys the activated neu-
rons are mirror neurons, this is not so in humans. In
humans, motor areas are endowed with two distinct popula-
tions of neurons: one merely sensory, the other merely
motor. The two populations do not communicate one with
another. Hence the mirror neurons do not exist in humans.
It was also claimed that, in order to prove “really” the
existence of mirror neurons in humans, one has to show
that they present the “repetition suppression”, that is that
they decrease their response not only following repetitive
observation of the same motor act done by another, but
123
234
also, crucially, following visual presentation of a motor act
after its execution. If mirror neurons do exist, they should
habituate in this cross-modal test.
As stressed by Logothetis (2008) there are serious diY-
culties in interpreting the results obtained with the repeti-
tion suppression technique. This is especially true for its
cross-modal variant. Repetitions suppression, in fact, takes
place when information arrives at a neuron using the same
or largely common pathways, but not when information
reaches a neuron using diVerent pathways. This because
repetition suppression is a phenomenon that occurs at the
synaptic level and not as a consequence of repetitive dis-
charge of a neuron (Sawamura et al. 2006).
It is important to keep in mind that, in the case of the
activation of mirror neurons, the communality of input in
the cross-modal test is typically lacking because during
action observation the input is coming mostly from STS,
while during voluntary movement the neurons are triggered
by commands arriving form the frontal lobe and other
higher order centers. The results of adaptation experiments
will depend therefore on the degree of input communality
to motor neuron that is introduced in the experimental
design. As one may expect from these considerations, the
results of the experiments using repetition suppression
technique produced contrasting results (e.g., Dinstein 2008;
Chong et al. 2008, Lingnau et al. 2009, Kilner et al. 2009).
However, the most recent data using this technique show
cross-modal adaptation and clearly prove the mirror neuron
existence (Kilner et al. 2009).
The proposed “two populations” hypothesis is also dis-
proved by experiments in which the authors tested whether
the same voxels were active in the mirror areas during
action observation and action execution (Gazzola and
Keysers 2009). This “shared voxels technique” showed that
the same voxels became active during both action observa-
tion and execution.
While the use of these new techniques may provide
interesting information on the organization the mirror net-
work, the neurophysiological reasoning underlying the
“two populations” hypothesis is very shaky. To believe that
the neurons located in motor areas that respond to action
observation are merely sensory or merely motor neurons
requires two assumptions. The Wrst is that, unlike monkeys,
human motor areas contain a large number of “displaced”
sensory neurons. The second is that these “displaced” neu-
rons do not communicate with motor neurons. While the
Wrst assumption is unlikely, but possible, the second is hard
to reconcile with all we know on the architecture of cere-
bral cortex.
Cerebral cortex is typically organized in columns with
rich connections between neurons in diVerent layers. To
postulate that information from STS reaching the parietal
and then motor areas remains in humans segregated from
123
Exp Brain Res (2010) 200:223–237
the motor output is absurd. First, even the staunchest
cognitivist would admit that sensory information is used
for motor purposes and not only for perception. Thus,
sensory information must reach cortical motor neurons.
Second, TMS studies (see above) showed that there is a
clear congruence between the observed motor act and the
activated motor representation. In other words higher
order sensory information describing motor acts activates
neurons that code those motor acts. But, if motor neurons
receive sensory information congruent with their motor
properties, these neurons are ‘mirror neurons’ by deWni-
tion. The “incommunicado populations” hypothesis
seems, therefore, a remote possibility rather than some-
thing grounded in physiological reality. However, even
wrong assumptions sometimes generate interesting exper-
iments.
Acknowledgments The study was supported by Fondazione Monte
Parma and by a grant (FIL) of the University of Parma to GR. M.F-D.
was supported by Fondazione Cassa di Risparmio di Ferrara. We thank
Rachel Wood for her comments on the article.
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