Transmodal Sensorimotor Networks during Action
Observation in Professional Pianists
B. Haslinger1, P. Erhard1, E. Altenmüller 2, U. Schroeder1,H. Boecker1, and A. O. Ceballos-Baumann1

Abstract& Audiovisual perception and imitation are essential formusical learning and skill acquisition. We compared professional pianists to musically naive controls with fMRI whileobserving piano playing finger–hand movements and serial
finger–thumb opposition movements both with and without
synchronous piano sound. Pianists showed stronger activations within a fronto-parieto-temporal network while observing
piano playing compared to controls and contrasted to perception of serial finger–thumb opposition movements.
Observation of silent piano playing additionally recruited

INTRODUCTION
Pianists spend years of extended and sustained training
beginning at the earliest possible age in childhood inorder to reach professional standard of musicianshipcomprising extreme upper-limb motor coordination
linked to auditory and sensorimotor feedback. In thisregard, musical skill acquisition shares many common
elements with the development of speech. In analogy
to infant learning of the native language, learning by listening, observation, and imitation of a teacher’s actions
are crucial steps in musical skill acquisition that are
widely applied in music pedagogics.
Musical training involves a strong functional association between motor performance and somatosensory
and auditory feedback. Professional musical training has
been demonstrated to induce adaptive functional andstructural changes within the motor system (Sluming
et al., 2002; Amunts, Schlaug, et al., 1997; Elbert, Pantev,
Wienbruch, Rockstroh, & Taub, 1995; Schlaug, Jancke,
Huang, Staiger, & Steinmetz, 1995) as well as within theauditory system (Schneider et al., 2002; Pantev et al.,
1998; Zatorre, Perry, Beckett, Westbury, & Evans, 1998;
Schlaug, Jancke, Huang, & Steinmetz, 1995). Sergent,
Zuck, Terriah, and MacDonald (1992) applied PET tostudy functional activations in musicians during musical sight-reading, active keyboard performance, and lis1

TU München, Munich, Germany, 2Institut für Musikphysiologie und Musiker-Medizin, Hochschule für Musik und Theater,
Hannover, Germany

D 2005 Massachusetts Institute of Technology

auditory areas in pianists. Perception of piano sounds
coupled with serial finger–thumb opposition movements
evoked increased activation within the sensorimotor network. This indicates specialization of multimodal auditory–
sensorimotor systems within a fronto-parieto-temporal network by professional musical training. Musical ‘‘language,’’which is acquired by observation and imitation, seems to betightly coupled to this network in accord with an observation–
execution system linking visual and auditory perception to
motor performance. &tening to the corresponding tones. They demonstrated afunctional association for these three domains within
the premotor, inferior frontal (BA 44), and parietal areas.
An EEG study suggested transmodal activation of thesensorimotor system by auditory input in professionalmusicians (Bangert, Haeusler, & Altenmuller, 2001).
Functional coupling of auditory and sensorimotor domains in musicians during passive observation of musicalmotor activity in association with corresponding acoustic input has not been studied with functional imaging
so far.
Previous functional imaging and neurophysiologicalstudies demonstrated involvement of a fronto-parietotemporal network during motor observation (Buccino
et al., 2001; Grezes & Decety, 2001; Perani et al., 2001;
Nishitani & Hari, 2000; Iacoboni, Woods, et al., 1999; Hariet al., 1998; Bonda, Petrides, Ostry, & Evans, 1996;
Grafton, Arbib, Fadiga, & Rizzolatti, 1996; Rizzolatti,
Fadiga, Matelli et al., 1996; Fadiga, Fogassi, Pavesi, &Rizzolatti, 1995; Decety, Pereni, et al., 1994). They
especially showed recruitment of inferior frontal areas
including parts of the inferior frontal gyrus (BA 44, 45)
and of the ventral premotor cortex (vPMC), of inferiorparietal/intraparietal areas as well as of the superior
temporal cortex (superior temporal sulcus [STS]) during
motor observation. Furthermore, activations within dorsal parts of the lateral premotor cortex (dPMC) as wellas within mesial premotor, prefrontal, and superior parietal cortices were associated with passive observationof movements.

Journal of Cognitive Neuroscience 17:2, pp. 282–293

Our fMRI study comparing professional pianists tomusically naive controls aimed at investigating theinfluence of musical training upon specialization of
the cortical motor observation network for musicalactivity and at studying transmodal auditory and sensorimotor activations. Observation of piano-related finger
movements was compared to perception of serial finger–
thumb opposition movements both with and without
piano sound. We expected stronger activations within
the fronto-parieto-temporal network in musicians duringobservation of meaningful piano-related movements
compared to observation of a resting hand and of serial
finger–thumb opposition movements. Furthermore, wehypothesized a transmodal recruitment of auditory association areas and of sensorimotor areas in pianists byperception of piano playing movements or piano tones,
respectively.

RESULTS
Differential Activation during Observation of
Meaningful Piano Playing Movements
We determined activation differences between musicians and musically naive controls when comparing observation of piano playing movements without soundto observation of a resting hand. This contrast was performed separately for right- and left-hand movements.
For observation of right-hand piano-related movements, professional musicians showed significantly
greater activation than controls within a cortical frontoparieto-temporal network: in the inferior frontal cortex
including the bilateral ventral parts of BA 6 (vPMC) andthe opercular part of the inferior frontal gyrus (BA 44),
as well as the bilateral inferior frontal gyrus, pars triangularis (BA 45), in the inferior parietal cortex (BA 40)(supramarginal gyrus) and the intraparietal sulcus (IPS)
bilaterally and in the bilateral STS. Additionally, thesuperior frontal cortex/dPMC (BA 6), mesial premotor
cortex/pre-SMA (BA 6), prefrontal cortex (BA 9/46),
orbito-frontal cortex (BA 47), postcentral gyrus (BA 2),
and superior parietal lobe (BA 7) were bilaterally stronger
activated in musicians. Interestingly, observation ofsilent musical motor activity evoked stronger signal
increases in pianists within the left primary auditory
cortex/transverse temporal gyrus (BA 41) and the left
superior temporal gyrus (BA 22, BA 42). Furthermore,musicians showed higher BOLD signal increases in thebilateral temporo-occipital cortex (BA 18/19/37), on thesubcortical level in the cerebellar hemispheres andstriatum bilaterally (Table 1, Figure 1).
For the comparison of observation of left-hand piano
playing movements without sound versus observation ofa resting hand between pianists and controls, a quite
similar activation pattern was found. Differing from theresults for right-hand movements, at the applied threshold, no activations were seen in the primary auditory

cortex and within the bilateral superior parietal cortex(Table 1).
To examine whether piano sound in association withpiano-related finger movements influences the activationdifferences, observation of piano playing movementswith sound versus observation of a resting hand wascontrasted between both groups. Professional musicians
showed significantly greater activation than controls
during observation of right-hand piano playing movements with piano sound in the left precentral cortex,
vPMC (BA 6) at the border to BA 44, inferior parietal
cortex (BA 40) (supramarginal gyrus) including a cluster
within IPS bilaterally, left superior frontal cortex/dPMC(BA 6), left prefrontal cortex (BA 9), left postcentral
cortex (BA 2), bilateral superior parietal cortex/precuneus (BA 7), right occipital cortex (BA 19), and both
cerebellar hemispheres (Table 2).
Intergroup comparison for observation of left-handpiano playing with sound versus observation of a resting
hand additionally showed higher activation in pianists
in the left inferior frontal gyrus (BA 44, 45), right inferior frontal gyrus (BA 45), and right dPMC with nodifferences in prefrontal, intraparietal, superior parietal,
occipital, and cerebellar areas (Table 2). Activation differences between both groups for observation of piano
playing movements with sound versus observation of a
resting hand for both hands were generally less extensive compared to the analysis of piano-related finger
movements without sound. Here no activation differences between both groups were detected within secondary auditory areas.

Differential Activation during Observation of
Piano Playing Compared to Serial Finger–Thumb
Opposition Movements
Comparing observation of meaningful piano playing toobservation of serial finger–thumb opposition movements without sound revealed activation differences
between pianists and controls in a subset of the areas
mentioned above suggesting their special role for the
training dependent meaning of the observed action:
for right-hand movements, the left inferior frontal cortex, namely, the vPMC (BA 6), opercular (BA 44) and prefrontal (BA 9) areas, right inferior parietal cortex (BA 40),
IPS bilaterally, bilateral dorsal premotor cortex/dPMC(BA 6), postcentral cortex (BA 2/3) bilaterally, left parietal operculum (BA 43), right caudate nucleus, and bilateral cerebellar hemispheres were activated stronger inmusicians (Table 3, Figure 2).
Activation changes in the intergroup comparison forleft-hand movements differed with respect to vPMC
activation that was stronger for musicians in the righthemisphere. No frontal opercular, prefrontal, intraparietal, and caudate signal differences were seen. Additionally, higher signal levels were detected for musicians

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Table 1. Fronto-Parieto-Temporal Areas with Stronger Activations in Musicians than in Controls during Observation of Piano
Playing without Sound Versus Observation of a Resting Hand
Without SoundArea

R Piano > Rest
ClusterBAxz

T levelCluster

BA

x

1

6

20

1

57

6.24*

1

6

22

Mesial premotor, pre-SMA

6

6

10

51

4.97*

6

R sup. frontal, dPMC

6

18

7

53

5.04*

L mid. frontal, vPMC

6

50

3

29

4.17*

L inf. frontal, pars opercularis

44

50

10

20

3.76*

45

55

16

3

2.91*

46

55

32

9

5.36*

51

29

2

44

55

12

L sup. frontal, dPMCy

L inf. frontal, pars triangularis

R inf. frontal

2

R inf. frontal, pars triangularis45yR inf. frontal, pars opercularisy

L Piano > Restz

T level

1

55

7.49*

2

11

57

6.49*

6

20

8

53

6.30*

2

6

51

4

31

5.49*

3

44

57

8

11

3.77*

45

55

16

5

5.14*

46

55

28

10

3.57*

4.07*

45

53

26

8

4.17*

3

2.90*

44

55

12

3

4.08*

4

y

R mid. frontal, vPMC

3

6

42

2

48

4.69*

5

6

59

2

37

4.64*

R mid. frontal

4

9

32

34

22

5.33*

6

46

50

46

18

3.63*

L mid. frontal

5

46

42

40

26

4.17*

7

46/10

40

40

24

3.66*

y

38

51

11

9

2.76

L sup. temporal

6

21

55

R sup. temporal, STS

7

22

53

19

1

4.47*

8

22

50

46

12

4.52*

40

61

38

24

3.72*

9

40

55

37

31

4.17*

42/22

59

23

9

4.53*

10

22

59

25

7

3.64*

L sup. temporal

42

59

26

14

4.20*

42

59

30

14

4.03*

L postcentral

2/40

53

25

38

4.49*

3

57

21

40

7.00*

L inf. parietal

40

65

24

29

3.63*

40

65

24

29

4.60*

R inf. parietal, supramarginal
L sup. temporal, STS

8

5 12

5.35*

L sup. temporal/inf. parietal

9

22/40

57

44

15

4.77*

22/40

59

38

20

5.45*

L transverse temporal gyrus

10

41

36

29

12

4.66*

41y

34

29

11

1.81

R inf. parietal/postcentral

11

40/2

46

31

46

4.37*

11

40

38

33

42

4.70*

R intraparietal, IPS

12

40

38

42

44

4.05*

12

40/7

44

46

56

4.14*

L intraparietal, IPS

13

40

32

39

35

4.64*

13

7/40

32

48

52

5.19*

R sup. parietal, precuneus

14

7

14

74

42

4.81*

7y

16

65

51

2.73

4.40*

y

12

61

53

2.83

L sup. parietal, precuneus

15

7

12

59

35

7

BA = Brodmann’s area; L = left; R = right. x, y, z express the position of the voxel with peak activation level ( p < .001, uncorrected, extent threshold
10 voxels) within the cluster in millimeters relative to the anterior commissure (AC) in the stereotactic space (Talairach & Tournoux, 1988).
yand italics indicate clusters that do not survive the threshold of p < .001, uncorrected, 10 voxels extent for the specific contrast. Here thehighest t value within the anatomically corresponding cluster is indicated. x = lateral distance from the midline (+ right,  left); y =anteroposterior distance from the AC (+ anterior,  posterior); z = height relative to the AC line (+ above,  below). dPMC, vPMC = dorsal,
ventral premotor cortex; SMA = supplementary motor area; STS = superior temporal sulcus; IPS = intraparietal sulcus; sup. = superior; inf. =inferior; mid. = middle.

*q < .05, FDR corrected.

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in the right superior parietal (BA 7) and left superior
temporal cortex (BA 22) (Table 3).
The direct comparison between pianists and controls
while observing finger tapping movements withoutsound versus rest did not show any significant differences. Nevertheless, activation differences were more
extensive and more significant for the contrast between
observation of musical motor activity versus the resting
condition compared to the contrast versus the nonmusical motor condition (Tables 1 and 3). We think that a‘‘resting’’ hand constitutes a relatively low-level baseline
condition compared to observation of finger tapping‘‘movements.’’ Accordingly, the direct comparison of observation of musical and nonmusical hand movements
analyzes more specifically the aspect of the meaning andcontent of musical motor activity that is modulated bymusical training in pianists.

Recruitment of the Sensorimotor System by
Auditory Input in Musicians
Only a small subset of areas was activated stronger inmusicians than controls when observing piano playing
with sound compared to serial finger–thumb opposition
movements with sound. For right-hand movements, theleft inferior parietal cortex (BA 40/2), the dPMC bilaterally, and the left superior parietal cortex (BA 7) wereactivated stronger in musicians; for left-hand movements, only the right superior parietal cortex (BA 5/7)
and left inferior parietal, somatosensory cortex (BA 1)
were activated stronger in musicians. The small extent
of those activation differences implies that in professional pianists, piano sound in association with serial
finger–thumb opposition movements could cause activation within the fronto-parieto-temporal association
network, therefore reducing activation differences between observation of piano-related and neutral movements with sound.
To further verify this aspect of activation of the sensorimotor system by auditory perception of piano soundin professional musicians, observation of serial finger–
thumb opposition movements with and without soundwas compared between both groups. We suppose thosemovements to have the same meaning and basic motor
input for both groups and therefore activation differences are suggested to be due to perception and computing of piano sounds. Here significant signal increases
were seen in pianists compared to controls in the left
vPMC and bilaterally in the cerebellar hemispheres. Additionally, a trend to significance at p < .01 was detectedin the inferior parietal cortex bilaterally, in the left parietal operculum, left superior parietal cortex, right prefrontal cortex, right superior temporal gyrus (BA 22),
cingulate cortex (BA 24), and occipital cortex (BA 18, 19).
For all conditions, on-line monitoring of the EMG
data during the test-run revealed no significant task-

related activity within the flexor muscles of both forearms in any subject.

DISCUSSION
In this study, we show that observation and recognitionof specific musical activity in professional pianists heavily
relies on a multimodal fronto-parieto-temporal network
including the inferior frontal gyrus (BA 44 and 45) andthe ventral part of the lateral premotor cortex (vPMC)
as well as the inferior parietal/intraparietal cortex andthe temporal cortex within the STS and the adjacenttemporal cortex (BA 21/22/42) bilaterally. Along with activation increases in dorsal fronto-parietal connections,this underlines the importance of this audiovisual–sensorimotor system for musical skills that are developed by sustained training and learning.
Whereas most studies of action observation have examined object-related actions, our study represents thefirst investigation involving action observation of piano
playing and finger–thumb opposition movements inprofessional pianists and musically naive controls. Inline with our results, inferior frontal opercular activationdifferences at the ventral border to the precentral cortex
in the context of motor observation are confirmed byrecent fMRI data (Grezes, Armony, Rowe, & Passingham,
2003; Hamzei et al., 2003). Despite substantial variability
in the anatomy of human BA 44 and 45 (Amunts,
Schleicher, et al., 1999), the stereotactic coordinates of
our inferior frontal activation clusters are in agreementwith previous data on activation patterns within thehuman inferior frontal cortex (Tomaiuolo et al., 1999).
A meta-analysis of PET studies on various motor observation tasks as well as a recent fMRI study referring tomean coordinates of previous imaging studies on motor
observation revealed comparable stereotactic localizations of activation within opercular parts of the inferior
frontal gyrus (BA 44), vPMC, middle/superior temporal
cortex, inferior parietal/intraparietal cortices, as well as
dorsal fronto-parietal areas (Grezes, Armony, et al., 2003;
Grezes & Decety, 2001).
Within this network, the polysensory STS could integrate information about movement recognition withspatial information for the analysis of the meaning andgoals of the observed behavior (Allison, Puce, & McCarthy, 2000; Oram & Perrett, 1996). This description
could be further processed within the inferior parietal/
intraparietal cortex, which has been suggested to represent an interface, integrating somatosensory and visual
information. STS and intraparietal areas have been described as heteromodal areas for cross-modal processing
and integration of audiovisual information (Calvert,
Hansen, Iversen, & Brammer, 2001; Calvert, Campbell,& Brammer, 2000). This analysis of the observed action
can be relayed to neurons with motor properties in theinferior frontal cortex. Functionally, those neurons are

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285

Figure 1. Areas showing significantly stronger activation in pianists than in controls during observation of right-hand piano playing
movements without sound compared to observation of a resting hand. Areas of activation ( p < .001, uncorrected) are projected onto a 3-D
surface rendering of a stereotactically normalized T1-weighted magnetic resonance image. The depth of the activated voxels relative to thebrain surface is coded in color intensity. The figure offers a view onto the right, left, and upper surface of the brain.

thought to describe an observed action in terms of its
goal (Iacoboni, Woods, et al., 1999; Gallese & Goldman,
1998; Jeannerod, Arbib, Rizzolatti, & Sakata, 1995).
Beside this inferior fronto-parieto-temporal system,
we also found a stronger involvement of dorsal frontoparietal connections in pianists compared to controls
during observation of musical hand actions. The superior parieto-dorsal premotor circuit has been suggested tobe responsible for motor control on the basis of somatosensory information (Rizzolatti, Luppino, & Matelli,1998). Superior parietal areas have been considered tointegrate multimodal somatosensory information of theperceived movement (Binkofski, Buccino, et al., 1999;
Grezes, Costes, & Decety, 1999; Iacoboni, Woods, et al.,1999). (Pre)SMA has been supposed to control theselection and preparation of motor programs that arecoded in the lateral parieto-frontal circuits. The prefrontal cortex provides inferior frontal areas with information
about the meaning and goal of an observed action(Rizzolatti & Luppino, 2001).
Bilateral activations of primary and secondary auditory
areas were detected in pianists compared to controls
when comparing observation of piano playing movements without sound to observation of a resting handand of serial finger–thumb opposition movements
without sound but not when opposing conditions withsound. This can be interpreted in the context of transmodal secondary auditory activation in musicians byvisual perception of piano playing due to the development of cross-modal processes as a result of long-termmusical training. Here, analogies can be establishedto the result of auditory activation by silent speech

Figure 2. Areas showing significantly stronger activation in pianists than in controls during observation of right-hand piano playing
movements without sound compared to observation of serial right-hand finger–thumb opposition movements without sound. Areas ofactivation ( p < .001, uncorrected) are projected onto a 3-D surface rendering of a stereotactically normalized T1-weighted magnetic
resonance image. The depth of the activated voxels relative to the brain surface is coded in color intensity. The figure offers a view onto
the right, left, and upper surface of the brain.

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Table 2. Fronto-Parieto-Temporal Areas with Stronger Activations in Musicians than in Controls during Observation of Piano
Playing with Sound Versus Observation of a Resting Hand
With SoundArea

R Piano > Rest

L Piano > Rest

Cluster

BA

x

y

z

T levelCluster

BA

x

y

z

T level

1

6

20

1

53

5.64*

1

6

20

1

55

5.47

R sup. frontal, dPMC

y

6

20

3

57

3.51

2

6

18

3

57

3.79

L inf. frontal, pars triangularis

45y

53

14

1

2.00

3

45

53

18

5

4.89

L inf. frontal, pars opercularis

y

44

57

12

3

2.65

44

51

12

5

4.24

6/44

63

2

11

5.05*

6

53

1

29

4.23

45y

53

31

6

2.83

45

50

25

4

3.95

9

59

15

27

2.72

40

67

33

29

3.93

40y

40

50

58

2.18

y

40

34

42

44

1.96

1

57

25

40

4.47

L sup. frontal, dPMC

L precentral, vPMC

2

R inf. frontal, pars triangularis
L mid./inf. frontal

3

9

40

27

26

5.00*

R inf. parietal, supramarginal

4

40

59

38

24

4.75*

R intraparietal, IPS

5

40

46

46

56

4.45*

4

y

5

L intraparietal, IPS

6

40

48

36

50

3.93

L postcentral, inf. parietal

7

2/40

65

20

25

4.06

R sup. parietal, precuneus

8

7

14

74

44

5.08*

7y

22

55

58

2.22

3.61

y

28

48

59

2.92

L sup. parietal

9

7

20

61

62

6

7

See notes to Table 1 for details.
*q < .05, FDR corrected.

perception as an example for cross-modal audiovisual
processing (Calvert, Bullmore, et al., 1997).
Additionally, these auditory activations and also thegreater activation differences in nonauditory corticalareas when contrasting observation of piano playing
without sound to observation of a resting hand and of
finger–thumb opposition movements without soundcould partly be due to musical (auditory) imaginationin our musicians. Aside from the temporal (auditory)
cortices, the lateral frontal, mesial frontal (SMA), andparietal areas are activated during the auditory imagination of music (Halpern & Zatorre, 1999; Zatorre,
Halpern, Perry, Meyer, & Evans, 1996). In contrast to previous data in nonmusicians showing a specialization of
the right temporal cortex for musical imagery (Halpern& Zatorre, 1999), our activations in musicians werelateralized to the left. As verbal processing during auditory imagery is thought to take place in the left superior
temporal cortex (Halpern & Zatorre, 1999), musiciansmight subconsciously attribute names of tones andintervals to the keys that are pressed by the observedhand. More likely, this shift from right hemisphericactivation to left in musicians compared to nonmusicians can be hypothesized as some kind of automatic
categorization, not necessarily verbal per se, that develops via musical training. The absence of auditory

imagery could in part explain comparatively smaller
activation differences when performing the same contrasts for observation of piano playing with sound.
Greater signal increases in pianists compared to controls within sensorimotor association areas due to theperception of piano tones synchronous to serial finger–
thumb opposition movements suggest a transmodal
recruitment of the sensorimotor system by auditory
perception of musical information in musicians. Thisis further supported by small activation differences forboth groups when comparing observation of meaningful piano playing movements with observation of serial
finger–thumb opposition movements both with sound.
It has previously been shown with EEG that after acertain time of musical practice already the isolated
perception of sounds without visual movement perception leads to activation within the premotor, primary
motor, parietal, and frontolateral areas that are also involved in actively playing a musical instrument (Bangert
et al., 2001). Neurons within the inferior frontal cortexof the monkey have been shown to discharge in response to action-related sounds (Kohler et al., 2002).
Learning-induced establishment of cross-modal links
to an auditory input has been demonstrated with
PET showing visual activation by isolated presentationof tones after a learning period with consistent pairing

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Table 3. Fronto-Parieto-Temporal Areas with Stronger Activations in Musicians than in Controls during Observation of Piano
Playing without Sound Versus Observation of Finger–Thumb Opposition Movements without Sound
Without SoundArea

R Piano > R FT
ClusterBAx

R sup. frontal, dPMC

1

6

22

L sup. frontal, dPMC

2

6

L inf. frontal

3

9

L inf. frontal, pars opercularisz

T levelCluster

BA

x

7

59

5.80*

1

6

22

24

3

61

5.38*

2

6

50

4

33

5.50*

59

44

y

L Piano > L FT

7

20

y

z

T level

9

62

4.27

22

3

55

4.43

9y

50

6

38

3.19

4.24

y

44

59

10

7

3.24

y

6

51

4

35

2.84

50

2

37

4.97*

6y

61

2

39

2.86

3

6

57

4

44

4.23

L sup. temporal

y

22

57

5

9

2.12

4

22

63

4

4

4.25

L sup. temporal

22y

59

40

15

3.01

5

22

63

42

19

4.09

2

38

23

38

4.47*

40y

40

34

51

2.85

40

38

29

35

3.78

3

59

14

27

4.27

2/40

59

23

45

4.25

3.69

y

43

63

11

14

2.99

y

L mid. frontal, vPMC

6

R mid. frontal, vPMC

R postcentral

4

R inf. parietal
L postcentral

5

L inf. parietal, operculum

63

43

13

19

6

R inf. parietal, IPS

6

40

34

48

43

4.18

40

36

44

52

2.96

L inf. parietal, IPS

7

40

36

41

39

3.81

40y

46

36

52

2.54

20

53

60

2.87

7

18

43

70

4.03

R sup. parietal

y

7

7

FT = fingertapping; see notes to Table 1 for further details.
*q < .05, FDR corrected.

of both modalities (McIntosh, Cabeza, & Lobaugh,1998). The auditory perception of piano tones might
therefore lead to a stronger recruitment of frontoparieto-temporal areas in pianists. This can explain
reduced between-group activation differences in the
motor observation network when comparing observation of piano playing and of serial finger–thumb opposition movements with sound.
Most of the fronto-parietal areas (apart from STS)
involved in observation have also been shown to be
activated to some extent during movement imagination(Binkofski, Amunts, et al., 2000; Grafton et al., 1996;
Stephan et al., 1995; Decety, Perani, et al., 1994).
Preparation of movement imitation (Krams, Rushworth,
Deiber, Frackowiak, & Passingham, 1998) and the intention to later imitate an action (Grezes, Costes, et al.,
1999; Decety, Grezes, et al., 1997) have been suggested
to modulate the recruitment of fronto-parietal areas
during action observation. A higher level of attention
associated with auditory imagery in conditions withoutsound or with motor imagery during perception of
piano sound can in part account for greater activation of

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the fronto-parieto-temporal network in pianists during
audiovisual perception of musical actions that havebeen practiced over many years. The action content andmeaning that is modulated by regular practice influences activation of inferior frontal, inferior parietal aswell as temporal motion areas. This is supported by previous data showing that especially observation of meaningful actions activates this cortical network (Decety,
Grezes, et al., 1997). The perceived piano-related movements and sounds are aligned with an internal motor
representation that has been established during many
years of musical motor training and analyzed withregard to their goal and meaning.
It is interesting to note that in our study activationdifferences within the fronto-parieto-temporal network
were generally more extensive and more significant
for observation of right-hand piano playing. The only
study that examined the effect of hand (right vs. left)on cerebral activation during motor observation demonstrated that during an observation–execution taskwith right hand motor responses, activation within the
STS was greater for passive observation of a right hand

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compared to observation of left-hand movements(Iacoboni, Koski, et al., 2001). The authors hypothesizedthat during observation STS activity ‘‘reflects an implicit
categorization of the moving hand as referred to thebody of the observer.’’ This contrasts a natural tendency
to imitate movements in a mirror configuration (righthand imitation of left-hand action). Furthermore, theright-handedness of all subjects in our study and predominance of the right hand in most piano compositions
may have contributed to stronger activation differences
between pianists and controls for observation of righthand piano playing movements.
It is tempting to speculate that some aspects of
the inferior fronto-parieto-temporal activation observedin the musicians reflect the operation of a ‘‘mirrormatching’’ system. Neurons with ‘‘mirror’’ properties (i.e.,
responding both to observation and execution of a
specific action) are found in the monkeys’ inferior
frontal area F5c (the homologue of human Broca’s
speech area) (Gallese, Fadiga, Fogassi, & Rizzolatti,
1996; Rizzolatti, Fadiga, Gallese, & Fogassi, 1996; diPellegrino, Fadiga, Fogassi, Gallese, & Rizzolatti, 1992),
within inferior parietal cortex (area PF) (Fogassi, Gallese, Fadiga, & Rizzolatti, 1998; Leinonen & Nyman,1979), as well as within the STS (Carey, Perrett, &Oram, 1997; Perrett et al., 1989). F5 neurons have also
been described to exhibit audiovisual properties responding to action-related sounds (Kohler et al., 2002).
This audiovisual observation–execution matching system is supposed to be involved in translating perception
to action by action recognition and understanding for
preparation of adequate motor responses and movement imitation (Umilta et al., 2001; Gallese & Goldman,
1998; Rizzolatti & Arbib, 1998). Long-term musical motor
learning and training by observation, listening, and imitation could support mirror-like neuronal computation.
The transmodal audiovisual network that we foundstronger activated in pianists during passive observationof musical motor activity could functionally couple
motor observation, auditory feedback, and imitation
for learning of musical skills. A specialized system seems
to develop during long-term piano training as an important element for the highly specific musical ‘‘intelligence’’ in professional musicians. This may constitutethe neurobiological basis of imitative, observative, andauditory-based teaching elements that are successfully
applied in music conservatories all over the world.

METHODS
Subjects
We investigated 12 professional pianists (6 women,
6 men, mean age 23.0 ± 2.4 years) and 12 musicallynaive control subjects (6 women, 6 men, mean age
25.0 ± 1.2 years). All subjects were strictly right handed
according to the diagnostic criteria of the Edinburgh

inventory (Oldfield, 1971). Musical professionalism wasdefined as having at least a completed conservatory
degree in piano playing. Mean history of piano practicewas 16.9 ± 2.7 years. Musically naive controls had tohave never played a musical instrument and not to havereceived any musical teaching beyond normal school
education. No subject had any history of neurologicaldisease. All subjects were naive with regard to the goalof the study. All subjects gave written informed consent
to the study in accordance with the guidelines fromthe Declaration of Helsinki. The study protocol wasapproved by the local Ethics Committee.
Task
The experimental task consisted in observation of different video sequences showing a real right or left hand
performing piano playing movements or serial finger–
thumb opposition movements with or without sound.
The video image was projected onto a transparent
screen using a video beamer. The scanning room wascompletely darkened. Subjects viewed the screen bymeans of a mirror that was centered above their eyes.
The video scenes were <108 wide and <7.58 high as
seen by the subjects with hand movements being centered to a range of <3.58 in width. The sound wastransmitted to the subjects via pneumatic earphones(EAR-Link 3a Cabot Safety, Indianapolis, IN, USA). Subjects were instructed only to watch the hand and to keep
their forearm, hand, or fingers completely relaxed. Botharms were kept supine, loosely enclosing a foam pad inorder to achieve a position that least resembles a natural
piano position. Five different conditions were presented
each for right- and left-hand movements respectively:First, sequences showed a right or left hand from abird’s eye view continuously performing silent monophonic piano playing movements pressing one key at a timewith a finger on a typical piano keyboard. Movements
were performed at a speed of 1 Hz within an interval of
10 tones, thereby avoiding extensive hand movements.
Finger movements were nonsequential, alternatingly involving all fingers of the hand and ‘‘randomly’’ distributed
over all keys within the interval (Figure 3A).
In a second condition, video sequences were shown
like in the first condition with the difference that thecorresponding piano sound was transmitted synchronous to the keypresses. Corresponding to the random
distribution of finger movements over the keys, thissound was no real melody or scale but a random
sequence of single tones. We chose this kind of random
and nonserial finger movements corresponding to nonmelodic random sequences of piano tones because wewere interested in the ‘‘basic’’ aspects of motor observation and sound perception. This avoids confounding
the results with ‘‘higher’’ aspects of musical perception
and musicality that cannot be compared between professional musicians and musically naive controls.

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Figure 3. Examples of the activation paradigm. The figure shows aright hand performing (A) ‘‘meaningful’’ piano playing movements ona keyboard and (B) ‘‘neutral’’ sequential finger to thumb opposition
movements supine in front of the keyboard. Both conditions were
presented with and without synchronous piano sound.

During the third condition, sequential finger to thumb
opposition movements with the right or left handsupine in front of the piano keyboard had to be observed (Figure 3B). These movements serially involved
finger to thumb oppositions of the index, middle, ring,and little finger in bidirectional order and were equally
performed at a frequency of 1 Hz without any sound.
The same sequential finger–thumb opposition movements were presented to the subjects as above in a
fourth condition where piano sound, representing random tonal sequences as in the piano conditions, wastransmitted synchronous to the finger movements. Piano tones appeared at a frequency of 1 Hz simultaneousto the time of contact between finger and thumb.
A right or left hand resting prone on the piano
keyboard was displayed as control condition.
Data Acquisition
fMRI measurements were performed on a 1.5 T Philips
Gyroscan NT scanner (Hamburg, Germany) upgraded to
Gyroscan Intera and equipped with a circular polarizedbirdcage head coil. A forehead restraining strip andvarious foam pads served for head fixation. The functional scans were acquired using a high-speed gradient-

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weighted EPI sequence (TR 3 sec, TE 50 msec, flip angle
908, FOV 230 mm, matrix 64  64 resulting in an in plane
resolution of 3.59 mm). A volume of 30 continuous axial
slices parallel to the AC–PC line with a single slice
thickness of 5 mm covering the whole brain was recorded during each scan. Additionally, a whole-brain 3-D
T1-weighted image was acquired for each subject for
high-resolution anatomical reference. The audio systemsound pressure level was optimized before each single
scanning session so that subjects could clearly hear thesound even during fMRI scanning.
We performed four block-design functional runs per
subject at 100 scans each with a single run duration of
300 sec. Two fixed experimental runs were designed foreach hand separately involving only observation of rightor left-hand movements, respectively. Each run wasgrouped into 10 epochs of 10 whole-brain EPI scans(30 sec). For the first run per hand, the five conditions
were alternated over epochs and repeated twice during
the run. The epoch order of the second run per hand
was derived from the first by mirroring the epoch order
before and after the fifth epoch. The same four experimental runs were presented for all subjects in pseudorandomized order, thereby always alternating left- andright-hand runs.
For controlling of eventual finger or hand movements
during motor observation, surface EMG electrodes wereattached over the forearm flexor muscles of both arms.
An additional experimental run without fMRI scanningwas performed at the end of the session for two-channel
surface EMG recordings. Subjects were naive about thegoal of this extra run.
The EMG amplifier as well as the video recorder andthe fMRI scanner were linked to a Lab-View based
electrophysiological monitoring system which allowed
the precise on-line registration of the EMG signal andthe fMRI timing parameters along with the experimental
paradigm (piano sound).
Data Analysis
Data were analyzed using SPM99 software (Wellcome
Department of Cognitive Neurology, www.fil.ion.ucl.
ac.uk/spm) based on the general linear model (Friston
et al., 1995). Calculations were performed on PCs running
LINUX and Matlab 5.3 (Mathworks, Sherborn, MA). The
first two scans of each run were omitted to allow equilibration of saturation effects resulting in a total of 98 scans
per run. Preprocessing of the data took several steps:
Images of each subject were realigned to the first image ofthe series, stereotactically normalized into a standard
space (Montreal Neurological Institute, MNI) approximating that of Talairach and Tournoux (1988) andsmoothed with a gaussian filter of 12  12  12 mm3.
A second-level random-effects approach was applied
for statistical analysis. On the first level, the four functional runs were entered into an individual design matrix

Volume 17, Number 2

for each subject. Here four different conditions per run,observation of right-hand piano playing or finger–thumb
opposition movements each with or without sound andaccordingly observation of left-hand piano playing or
finger–thumb opposition movements each with or without sound, were defined explicitly with the control
condition modeled implicitly. Data were convolved witha canonical hemodynamic response function. A highpass filter was applied for filtering low-frequency noise(range of cutoffs: 240–264 sec). Four different contrasts
were defined comparing observation of piano playing
movements (Piano) without or with piano sound (Silent/Sound) to observation of sequential finger–thumb opposition movements (Sequence) and to observation of a
resting hand (Rest) separately for right- and left-handmovements, respectively. Additionally, to detect if perception of piano sound in association with serial finger–
thumb opposition movements leads to an increase insensorimotor activation, we compared observation of
finger–thumb opposition movements with sound (bothright and left hand together) to observation of finger–
thumb opposition movements without sound. This resulted in a total of nine contrast images per subject:
1. Right Piano Silent–Right Rest
2. Right Piano Sound–Right Rest
3. Right Piano Silent–Right Sequence Silent
4. Right Piano Sound–Right Sequence Sound
5. Left Piano Silent–Left Rest
6. Left Piano Sound–Left Rest
7. Left Piano Silent–Left Sequence Silent
8. Left Piano Sound–Left Sequence Sound
9. Right + Left Sequence Sound–Right + Left
Sequence Silent
The corresponding contrast images were enteredinto a second level (random effects) analysis for groupcomparison. A two-sample t test was applied for calculating activation differences between the group of professional pianists and the group of musical laymen.
Based on the results of previous functional imaging
studies on motor observation and our a priori hypotheses, a statistical threshold of p < .001, uncorrected(10 voxels extent threshold), was considered to show
significant activation. To further corroborate the findingsa threshold of p < .05 FDR corrected was applied to thedata (Genovese, Lazar, & Nichols, 2002). For localization
within the stereotactic space of Talairach (Talairach &Tournoux, 1988), the coordinates of activated voxels
were transformed from MNI to Talairach coordinates
using a routine proposed by M. Brett (www.mrc-cbu.
cam.ac.uk/Imaging/mnispace.html).

Acknowledgments
This work was supported by the SFB 462 ‘‘Sensomotorik,’’
Deutsche Forschungsgemeinschaft Bonn and the ‘‘Gerhard
und Irmgard Schulz Fond,’’ Neurologische Klinik, Klinikum

Rechts der Isar. We thank all subjects, especially theprofessional pianists, for participating in the study. Special
thanks to Martina Bauer (‘‘Hochschule für Musik und
Theater,’’ Munich) and Dr. Thomas Hitzlberger (‘‘Richard
Strauss Konservatorium,’’ Munich) for their support in recruiting the pianists.
Reprint requests should be sent to Bernhard Haslinger,
Neurologische Klinik und Poliklinik, TU-München Möhlstrabe
28 D-81675 Munich, or via e-mail: b.haslinger@lrz.tumuenchen.de.
The data reported in this experiment have been depositedin the fMRI Data Center (www.fmridc.org). The accession
number is 2-2004-1177X.

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