Brain Topogr (2008) 21:9399

DOI 10.1007/s10548-008-0066-1

ORIGINAL PAPER

EEG Correlates of Action Observation in Humans

Elisa Mira Holz Michael Doppelmayr
Wolfgang Klimesch Paul Sauseng

Accepted: 26 August 2008 / Published online: 9 September 2008

Springer Science+Business Media, LLC 2008

Abstract To investigate electrophysiological correlates

of action observation electroencephalogram (EEG) was
recorded while participants observed repetitive biological
(human) or non-biological movements (at a rate of 2 Hz).
Steady-state evoked potentials were analyzed and their
neural sources were investigated using low resolution
electromagnetic tomography analysis (LORETA). Results
revealed significantly higher activation in the primary
motor and premotor cortex, supplementary motor area as
well as the posterior parietal cortices during observation of
biological movements, supporting mirror properties of
cortical motor neurons. In addition, interregional communication was analyzed. Increased coherence for distributed
networks at delta (0.54 Hz) and lower alpha (810 Hz)
frequencies were obtained suggesting integration and
functional coupling between the activated cortical regions
during human action observation.
Keywords Alpha
Coherence
Delta
LORETA

Mirror neuron system
Oscillations

Introduction
To facilitate understanding and imitation of other persons
behaviour a system matching observed actions with ones
E. M. Holz
M. Doppelmayr
W. Klimesch
P. Sauseng (&)
Department of Psychology, University of Salzburg,
Hellbrunnerstr. 34, 5020 Salzburg, Austria
e-mail: paul.sauseng@sbg.ac.at
P. Sauseng
Department of Neurology, University Hospital Eppendorf,
University of Hamburg, Martinistr. 52, 20246 Hamburg,
Germany

own motor representations is implemented in the brain.

Brain imaging studies postulate that this action observationexecution matching system is associated with a frontoparietal network, in particular the inferior frontal gyrus, the
inferior parietal lobe and the premotor cortex (Gallese et al.
2004; Rizzolatti and Craighero 2004; Iacoboni et al. 2005).
This system, also known as Mirror Neuron System (MNS),
enables us to understand what others are doing by integrating the observed movement into an internal simulation
of this action (Oberman et al. 2005; Rizzolatti and Craighero 2004). This kind of mental imitation of the observed
action seems to be necessary for imitating others behaviour
(Gallese and Goldman 1998; Agnew et al. 2007). Interestingly reaction times (RT) during imitation are shorter when
finger movements have to be imitated than when non-biological ones are presented (Brass et al. 2000; Kessler et al.
2006). Buccino et al. (2004) showed that actions that
belong to the observers own motor system or are similar to
them but from other species are mapped on the observers
own motor system. But actions that do not belong to it are
predominantly analyzed on their visual properties. It seems
as if cortical motor neurons or neurons with motor functions
belong to a MNS which preferentially processes human
actions whereas non-biological motions or motions of
objects are rather processed by the visual system.
Motor neuron activity is characterized by oscillatory
changes in the 812 Hz range (for a review see Pineda
2005). 812 Hz activity over sensorimotor areas is suppressed during execution (Manganotti et al. 1998) and
imagination of human actions (Pfurtscheller and Neuper
1997; Pfurtscheller et al. 1999; Neuper et al. 2005) but also
during observation of human movements (Hari et al. 1998;
Cochin et al. 1999; Rossi et al. 2002; Muthukumaraswamy
and Johnson 2004). This latter phenomenon was considered to reflect the activity of mirror neurons. EEG studies

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comparing brain activity during observation of biological

and non-biological motion showed that activity around
10 Hz over central (sensorimotor) areas is significantly
more suppressed during the observation of biological
(human) movements than observation of scrambled motion
(Ulloa and Pineda 2007) or motion of objects (Oberman
et al. 2005). Despite many findings supporting the existence of a MNS tuned for human movements the results
remain discordant. For instance also visuomotor priming
for robotic movements (Press et al. 2005) and activation in
MNS caused by robotic movements was found (Gazzola
et al. 2007; Oberman et al. 2007).
For visual stimulation most of the above cited EEG
studies used continuous motion videos without a clear
stimulus onset. These ongoing tasks make it hard to analyse event-related potentials. Therefore, these EEG studies
only relied on reactivity of certain EEG frequency bands as
indicator for mirror neuron activity. Here, steady-state finger or object movements at a rate of 2 Hz were presented.
EEG activity phase-locked to the observation of rhythmical
down-movements of the finger or an object was analyzed.
The steady-state evoked potentials elicited by this kind of
stimulation in the present study were then used to localize
mirror neuron activity in 3-D source space. Oscillatory
brain activity (amplitude estimates as well as interregional
synchronization) has turned out to play a major role during
various motor functions (Pfurtscheller and Neuper 1997;
Manganotti et al. 1998; Neuper et al. 2005; Pineda 2005).
However there is a lack of research on these brain parameters regarding the observation of biological and nonbiological movements. Therefore, a further aim was to
examine what kind of information about mirror neuron
activity can arise from task related coherence between brain
regionspredominantly in the frequency range around
10 Hzwith regard to the MNS. Subjects viewed kinematically matched rhythmical non-biological and biological
movements. With regard to above cited findings we
expected higher activity in terms of steady-state evoked
potential amplitude in fronto-central and parietal regions for
biological motion. We also hypothesized higher interregional synchronization between the activated brain regions
in particular at the EEG alpha frequency band for observation of biological actions (Manganotti et al. 1998).

Methods
Forty-five subjects participated voluntarily in this study.
Four subjects were excluded from analysis because of
muscle or eye-blink artifacts. The sample of 41 subjects (15
men and 26 women) were at the average age of 23.41 years
(SD = 2.91). All participants were right-handed, had normal or corrected-to-normal vision and had no history of

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Brain Topogr (2008) 21:9399

neurological disorders. They all gave informed consent

according to the Declaration of Helsinki and were naive
with respect to the purpose of the experiment.
EEG was recorded in an auditory shielded room during
3 conditions. In the moving finger condition (biological
movements) the subjects observed movies of fast repetitive
(human) finger movements (mouse clicks with the index
finger of the right hand onto the left button of a computer
mouse) at a rate of 2 Hz (visually paced). In the moving
object condition (non-biological movements) subjects
viewed the same movements generated by a metal bar
instead of a finger. To ensure that attention was paid to the
videos participants had to absolve a kind of continuous
performance task. Within the blocks of either finger or bar
motions there were button presses to the right instead of the
left mouse button, three to five times for each condition.
These events had to be detected and silently counted (later
reports suggest a 100% correct performance in all subjects). The paradigm was organized as block design with
2 min finger movements and 2 min bar movements
(resulting in 240 movements for each condition). Half of
the subjects first viewed the moving finger condition and
then the moving object condition and half of the subjects
viewed the conditions in the inverse order.
To compare the data from the moving finger and moving
object condition also to a neutral condition, EEG recordings during a 2 min baseline measure (resting condition)
were used as an additional control condition. In this condition subjects were required only to keep their eyes open
and fixate the middle of a computer screen without any
additional task or stimulation.
EEG was recorded using a BrainAmp amplifier (Brain
Products, Germany) with 32 channels and a sampling frequency of 500 Hz. Twenty-seven AgAgCl electrodes were
placed according to the extended 1020 system at the
positions Fp1, Fp2, F7, F3, Fz, F4, F8, FT7, FC3, FCz, FC4,
FT8, T3, C3, Cz, C4, T4, Cp3, CPz, Cp4, T5, P3, Pz, P4, T6,
O1, O2 and were recorded against a common reference at
the nose. In addition, the vertical electrooculogram (EOG)
was recorded. To control for minimal finger movements we
recorded electromyogram (EMG) at the finger flexors of the
right hand. Impedance was kept below 15 kX.
Data analysis was done using Vision Analyzer (Brain
Products, Germany). First EEG-signals were offline re-referenced to digitally linked earlobes. Ocular correction as
suggested by Gratton et al. (1983) was applied. Remaining
ocular and muscular artefacts were manually eliminated. A
low-cutoff filter of 0.5 Hz and a high-cutoff filter of 50 Hz
with a notch filter at 50 Hz were used (EMG was recorded
between 10 and 250 Hz with a notch filter at 50 Hz). To
control for the possibility that differences at EEG activity
over sensorimotor regions between conditions are elicited
by minimal finger movements, EMG was compared between

Brain Topogr (2008) 21:9399

95

the two types of tasks. Therefore the EMG was rectified and
the average activity of each block was statistically compared
between conditions using paired sample t-test. The t-test was
not significant, t (40) = .45, P = .66. This shows that EEG
differences between conditions cannot be explained by
artificial finger movements. This does not exclude the possibility that subjects performed small finger movements
during the experiment; however the non-significance of the
statistical comparison between conditions demonstrates that
subjects did not move fingers differently during the moving
finger and the moving object condition.
EEG data were segmented into 500 ms epochs, each
representing observation of one complete movement (at a
rate of 2 Hz). For the resting condition, data were also
segmented into epochs of 500 ms.
Segments were then averaged separately for the moving
finger and the moving object condition. On average the
number of artefact free trials was 208.37 and 205.85 for the
two conditions, respectively. Averaging of trials resulted in

F3

Fz

FC3

FCz

F4

steady-state evoked potentials (SSEPs; Gerloff et al. 1997)

which are shown in Fig. 1a. For the resting condition no
SSEPs could be calculated as there was no rhythmical
visual stimulation in this condition.
Positive and negative maxima in the SSEP were determined using peak detection. Then the difference between
the positive and negative peak amplitude was calculated.
This was done separately for the moving finger and the
moving object condition. Then these values were entered
into non-parametrical comparisons (Wilcoxon tests)
between the two conditions separately for each recording
site.
SSEPs were used for source analysis. Therefore, low
resolution electromagnetic tomography analysis (LORETA;
Pascual-Marqui et al. 1994) was performed. The current
source density (CSD) was calculated for each time frame of
the 500 ms analysis interval of the SSEPs for 2,394 cortical
voxels, under the assumption that adjacent voxels have
similar activity (Pascual-Marqui et al. 1994). Then all time

[ms]

FC4

C3

Cz

C4

CPz

CP4

CP3

P3

Pz

P4

A
O1

O2

5 V
0
- 5 V
-250 ms

0 ms

250 ms

Fig. 1 Steady-state evoked potentials elicited by action observation

at scalp and source level. (a) Steady-state evoked potentials for the
moving finger (black) and the moving object condition (red line) for
selected electrodes. Brain potentials are phase-locked to the down
movements (at 0 ms) of the observed finger or bar motion. (b) Low
resolution electromagnetic tomography analysis (LORETA) was used
to localize the neural sources underlying the difference between the
two conditions on the electrode level. Red colour indicates higher

current source density (CSD) in the moving finger compared to the

moving object condition (P \ .01; A = anterior, P = posterior,
L = left, R = right). Note that there is higher activity in bilateral
primary motor cortices (M1), premotor cortices, supplementary motor
areas, right prefrontal cortex and superior and posterior parietal
cortices for the moving finger compared to the moving object
condition

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frames were averaged for each condition separately. CSD for

every voxel was compared between both conditions running
voxel-wise non-parametrical statistics as implemented
in LORETA on the 1% significance level (corrected for
multiple comparisons; non-parametric bootstrapping was
used for significance testing, for details see Nichols and
Holmes 2002).
To compute EEG coherence we applied Fast Fourier
Transformation (FFT) to the 500 ms epochs (for the moving
finger, moving object and resting condition separately) and
calculated coherences of all combinations of channels
(n = 351) for the frequency bands: delta (0.54 Hz), theta
(48 Hz), lower alpha (810 Hz), upper alpha (1012 Hz)
and beta 1 (1220 Hz), beta 2 (2030 Hz) and gamma (30
50 Hz). Previous research suggests these frequency bands
to play an important role in cognitive and motoric processes
(Pfurtscheller and Neuper 1997; Gerloff et al. 1998; Neuper
et al. 2005; Pineda 2005; Calmels et al. 2006). Coherence is
a measure of signal similarity between different electrode
sites. Values can range between 0 and 1where 0 indicates
no similarity and 1 stands for absolute similarity. Usually
coherence values are not normally distributed, thus values
were Fisher-z-transformed. Wilcoxons signed rank tests
comparing values between conditions on the 1% significance level were run for each electrode pair (Rappelsberger
and Petsche 1988). To examine whether there was a
coherent pattern of either increase or decrease of coherence
in the moving finger condition compared to the moving
object condition or between these conditions and the resting
condition the number of electrode pairs with a significant
increase of coherence were compared with the number of
electrode pairs with significant decrease of coherence using
chi-square tests. Therefore, if there was neither a clear
pattern of increase nor a clear pattern of decrease of
coherence over electrode pairs, i.e. when there was an equal
distribution between electrode pairs showing stronger
coherence in the moving finger condition and electrode
pairs with higher coherence in the moving object or resting
condition, respectively, this was indicated by non-significance of the chi-square test. The chi-square testing was
done separately for each of the 7 frequency bands and each
comparison (moving finger vs. moving object, moving finger
vs. resting condition and moving object vs. resting condition). To correct for this multiple testing Bonferroni
correction was applied to the results.

Results
SSEPs on the Scalp Level
Wilcoxon paired comparisons indicate that there were
SSEPs with larger amplitude in the moving finger than the

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Brain Topogr (2008) 21:9399

moving object condition at the following recording sites:

F3, Fz, F4, FC3, FCz, FC4, Cz, C4, CPz, CP4, O2 (all
Z [ 3.12, P \ .05, Bonferroni corrected).
Current Source Density of SSEPs
LORETA results indicate that there was higher bilateral
activation in the moving finger condition in the premotor
cortex, supplementary motor area (SMA), primary motor
cortex as well as the posterior parietal cortex (PPC),
whereas the latter activation was predominant in the right
hemisphere. In addition higher activation was observed in
the right prefrontal cortex (t = 3.89, P \ .01). As can be
seen in Fig. 1b there was no single voxel were the moving
object condition showed higher CSD than the moving finger condition.
Interregional Coherence
Moving Finger Versus Moving Object Condition
Coherence analysis revealed stronger synchronization in
the moving finger as compared with the moving object
condition for lower alpha (v2 = 22.27, P \ .01) and delta
(v2 = 16.95, P \ .01) frequency bands, as depicted in
Fig. 2. In the lower alpha frequency band a fronto-parietal
network including recording sites overlying the premotor
cortex and primary sensorimotor cortex of the left (FC3,
C3, CP3) and the right (FC4, C4, CP4) hemisphere and the
frontocentral cortex including the SMA (Fz, FCz, Cz)
(Homan et al. 1987) were coherently active during observation of finger movements. Delta coherence was found
between occipital and central electrode sites with predominant coupling between the right primary visual cortex
(O2) and the sensorimotor cortex and the SMA. Higher
coherence in the moving object condition was found for the
theta frequency band (v2 = 99.56, P \ .01), extending
over a wider fronto-parietooccipital network. No systematic differences in coherence between conditions were
found for upper alpha, beta 1, beta 2 and gamma.
Moving Finger Versus Resting Condition
Stronger interregional synchronization was found for delta
(v2 = 137, P \ .01), theta (v2 = 11. 94, P \ .01), lower
alpha (v2 = 77.76, P \ .01) and gamma (v2 = 27.22,
P \ .01) in the moving finger condition than during rest.
Delta coupling includes a wide fronto-parietooccipital
network (see Fig. 2). In the theta frequency higher synchronization in a fronto-parietal network was found for the
moving finger condition than for rest. For lower alpha
higher coherent coupling was found between electrode
sites overlying the bilateral sensorimotor and premotor

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97

Fig. 2 Coherence at delta, theta

and lower alpha frequency
bands. Red connections denote
higher coherence during
observation of finger movements
(P \ .01) compared to moving
object (a) or resting condition
(b) and for moving object
compared to a resting condition
(c). Blue connections indicate
higher coherences during the
moving object or resting
condition (a and b, respectively)
than during observation of the
moving finger condition, or in
(c) higher coherence in the
resting condition compared to
the moving object condition in
delta, theta and lower alpha
frequency bands

cortex, the supplementary motor area (SMA), the posterior

parietal cortex (PPC) and the right temporal cortex. Within
Gamma frequency left temporal sites are coupled with
central leads. For Beta 1 we found higher synchronization
for the resting condition within a broad distributed network
(v2 = 41.88, P \ .01). No significant effects were found
for upper alpha and beta 2.
Moving Object Versus Resting Condition
We found a higher degree of synchronized activity in the
moving object condition for delta (v2 = 22, P \ .01) and
theta (v2 = 51.02, P \ .01). The coupling for delta is spread
over frontal and parietal regions. For theta global coupling
was found. Stronger desynchronization in the moving object
condition than for rest were found for beta 1 (v2 = 65.09,
P \ .01) and beta 2 (v2 = 40.16, P \ .01), extending over a
wide distributed global network. There were no significant
results for lower alpha, upper alpha and gamma.

Discussion
Brain activity evaluated by the means of LORETA during
observation of biological versus non-biological movements
was assessed. The relative brain activity should show the
location of mirror neurons. The results indicate that regions

are stronger activated during observation of biological

versus non-biological movements, in particular the premotor cortex, supplementary motor cortex, primary motor
cortex, and the posterior parietal cortex (PPC). This is well
in line with other studies showing that premotor, supplementary and primary motor regions contain neurons which
have mirror neuron properties (Hari et al. 1998; Cochin
et al. 1999; Rossi et al. 2002; Perani et al. 2001; Muthukumaraswamy and Johnson 2004; Oberman et al. 2005;
Ulloa and Pineda 2007).
There is agreement that the PPC is responsible for visuomotoric integration (Buneo and Andersen 2006;
Iacoboni 2006). However the right posterior parietal activation seems also to be related to self-recognition during
imitation whereas the left posterior parietal activity is
related to tool use (Iacoboni 2006). The higher right posterior parietal activation during observation of biological
movements as found in this experiment might reflect mirror
neuron activity evoked by the recognition of the human
hand. Thus, the posterior parietal cortex seems to be an
important link between visuomotoric integration and the
emergence of an internal representation of an action.
Although there are a lot of findings describing the
localization of the MNS, little is known about the interaction within this network and with other regions. We
found interregional coupling predominately at lower alpha
and delta frequency during the observation of finger

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movements. Coherence within the EEG lower alpha frequency range (810 Hz) showed a similar distribution
compared with coherence during execution of repetitive
finger movements in other studies, predominantly over
fronto-central brain regions (Manganotti et al. 1998).
However, it should be noted that in the present experiment
subjects only observed movements but did not perform the
observed actions. Therefore, it cannot derive from the
present data whether during the imitation of actions there
would be identical neural activity as during their
observation.
Several studies have shown that alpha is the prevailing
frequency of coupling during repetitive finger movements
(Manganotti et al. 1998; Pollok et al. 2005; Toma et al.
2002). These results are in line with findings of Calmels
et al. (2006). They found alpha synchronization over
fronto-central brain regions during both action execution
and observation. The comparison of the oscillatory brain
activity during observation of biological and non-biological motion makes obvious that this specialized 810 Hz
network underlies predominately the observation of finger
movements. This is further evidence for mirror neurons,
which seem to be tuned for observation of biological
actions. Motor-relevant regions were in addition coupled to
the PPC at lower alpha frequency in the present study. This
confirms the assumption that the PPC is functionally linked
to the MNS. These findings are also underpinned by the
fact that there was no similar pattern for the non-biological
movement condition (compared with a resting condition).
Additionally, we found global synchronization within
delta (0.54 Hz) frequency during observation of finger
movements whereas there was less coupling for the
observation of movements of an object. Coupling within
delta seems to be a mechanism to integrate the rhythmic
visuo-motoric information in terms of synchronization
between parietal and central regions. Findings confirm that
lower frequencies are related to global binding which
means synchronization of a large neuronal network (von
Stein and Sarntheim 2000). This effect might be reinforced
by the 2 Hz rhythmic stimulation of the experimental
design (which is indicated by the fact that there is an even
stronger effect when the moving finger condition was
compared to the resting condition without rhythmical
stimulation). As described above mirror neuron activity in
the PPC and coupling between PPC and motor cortices
may affect the visuo-motoric integration of biological
movements. Thus we suggest that these processes are
crucial for encoding of the properties (i.e. dynamics) of
action and therefore enable imitation. With other words,
these processes seem to facilitate imitation to biological
action cues. Indeed findings indicate that during imitation
reaction times and the degree of synchronization between
PPC and the premotor cortices are correlated in an early

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time window after presentation of a cue (Kessler et al.

2006). This was interpreted as a behavioural advantage for
biological cues. Thus, besides the understanding of the
intentions of others the understanding of the dynamics of
movements can be seen as necessary for imitation of an
action (Wolfensteller et al. 2007).
We found a higher number of coherent electrode pairs in
the theta band for fronto-parietal long range connections
for the moving object than in the moving finger condition
(see Fig. 2). Sauseng et al. (2007) showed that motor tasks
requiring a higher degree of executive control exhibit
stronger fronto-parietal theta coupling. As the observation
of biological movements is more familiar than non-biological motions it is supposed that the latter requires more
executive control and thus elicits stronger theta coupling
than observation of biological movements. Compared to
the resting condition both the biological and the non-biological movement condition showed a fronto-central
increase of theta coupling and fronto-posterior decrease of
theta coherence. The synchronization in the non-biological
movement condition was stronger, underpinning the
interpretation that this condition requires more executive
functions. At beta 1 there was decreased coherence in the
biological and the non-biological movement conditions
compared to the resting condition. This effect did not differentiate between the movement observation conditions
and thus seems to be rather unspecific and related to visual
stimulation. For gamma there was stronger interregional
coupling in the moving finger condition and for beta 2
decrease of coherence compared to the resting condition,
which similarly to beta 1 did not dissociate between biological and non-biological movement conditions.
Concluding, this study replicated, on the basis of SSEPs,
the findings indicating that a distributed bilateral network
including primary motor cortex, supplementary motor area,
premotor cortex and (right) parietal cortex is selectively
active during observation of biological movements and less
during observation of non-biological motion. These results
provide mirror properties of these regions; however the
lack of an execution condition in the present study limits
evidence for an action observationexecution matching
system. We could show that these fronto-central regions
are coherently active while biological but not non-biological movements were observed, in particular at lower alpha
frequency. As brain activity in the alpha frequency range is
related to motoric activation we conclude that this is a
further indicator for mirror neuron activity.
Acknowledgements This research was supported by the Austrian
Academy of Sciences P_145001_1. PS is recipient of an APART
fellowship by the Austrian Academy of Sciences. Thanks also to
Sieglinde Gruber and Kerstin Hoedlmoser and several undergraduate
students at the University of Salzburg who helped with data
acquisition.

Brain Topogr (2008) 21:9399

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