MEDICINE, MUSIC AND THE MIND

Neurology of musical performance

Eckart Altenmller

Eckart
Altenmller
Director, Institute of
Music Physiology
and Musicians
Medicine,
University of Music
and Drama
Hannover, Germany
Clin Med
2008;8:41013

ABSTRACT Performing music at a professional

level requires the integration of multimodal sensory and motor information and precise monitoring of the performance via auditory feedback.
In the context of Western classical music, musicians are forced to reproduce highly controlled
movements almost perfectly with a high reliability.
These specialised sensorimotor skills are acquired
during extensive training periods over many years.
The superior skills of musicians are mirrored in
functional and structural plastic adaptations of
sensorimotor and auditory systems of the brain.
Auditory-sensorimotor integration, for example, is
accompanied by rapid modulations of neuronal
connectivity in the time range of 20 minutes.
Finally, dysfunctional plasticity in musicians,
known as musicians dystonia, leads to deterioration of extensively trained fine motor skills.
Musicians dystonia may be caused by training
induced dysplasticity with pathological fusion of
central nervous representations in sensorimotor
cortical and subcortical brain regions.
KEY WORDS: brain plasticity, music performance,
expertise, sensorimotor integration, musicians
dystonia

Apollos gift: music making as a stimulus

for brain plasticity
Performing music at a professional level is one of
the most complex human accomplishments. Music,
as a sensory stimulus, and is structured along several
dimensions. Moreover, making music requires the
integration of multimodal sensory and motor
information and precise monitoring of the motor
performance via auditory feedback. In the context of
Western classical music, musicians are forced to
reproduce highly controlled movements almost
perfectly and with a high reliability. These specialised
sensorimotor skills require extensive training over
many years, starting in early infancy and passing
through stages of increasing physical and strategic
complexities.
Sensorimotor skills in musicians are usually automated by many repetitions whereas aural skills are
typically refined through a broad variety of listening
experiences. Both types of skills are not represented
in isolated brain areas, however, but rather depend
410

on the multiple connections and interactions established during training within and between the different regions of the brain. The general ability of the
central nervous system (CNS) to adapt to changing
environmental conditions and newly imposed tasks
during its entire lifespan is referred to as plasticity.
In music, learning through experience and
training is accompanied by remarkable plastic adaptations, which are not only reflected in modifications
of the brains neuronal networks, as a result of a
strengthening of neuronal connections, but also in its
overall gross structure. It is known, for example, that
music practice enhances myelination, grey matter
growth and fibre formation of brain structures
involved in the specific musical task.1 There are two
main reasons why researchers believe that these
effects on brain plasticity are more pronounced in
instrumental music performers than in other skilled
activities. First, musical training usually starts very
early, sometimes before age six, when the adaptability of the CNS is highest, and second, musical
activities are strongly linked to positive emotions,
which are known to enhance plastic adaptations.
This paper will focus on new insights concerning
brain mechanisms involved in musical performance
and practice. Changes in brain networks and structures accompanying musical achievements, and the
neural foundations of training strategies such as
mental and observative practice will be discussed.
The paper will conclude with the effects related to
maladaptive changes of brain networks, resulting in
movement disorders such as musicians dystonia.

Wiring the brain: music making as a

sensorimotor integration task
Performance-based music making relies primarily on
a highly developed auditory-motor integration
capacity, which can be compared to the phonological
loop in speech production. In addition, somatosensory feedback constitutes another basis of high
level performance. Here, the kinesthetic sense, which
allows for control and feedback of muscle and tendontension as well as joint positions which enable continuous monitoring of finger, hand or lip position in the
frames of body and instrument coordinates (eg the
keyboard, the mouthpiece), is especially important.
One special quality of musicianship is the strong
coupling of sensorimotor and auditory processing.
Clinical Medicine Vol 8 No 4 August 2008
Royal College of Physicians, 2008. All rights reserved.

Neurology of musical performance

Practising an instrument involves assembling, storing and constantly improving complex sensorimotor programs through
prolonged and repeated execution of motor patterns under the
controlled monitoring of the auditory system. Many professional pianists, report that their fingers move more or less
automatically when they are listening to piano music played by
a colleague. In a cross-sectional experiment, strong linkages
between auditory and sensorimotor cortical regions develop as
a result of many years of practice.2 Using functional magnetic
resonance imaging professional pianists were asked to listen to
simple piano tunes without moving their fingers or any other
body part. Figure 1 demonstrates the increase in activation of
professional pianists in comparison to non-musicians. There is
an impressive activation of the motor cortex demonstrating the
sub-conscious or automated auditory-motor co-activation.
Furthermore, in a longitudinal study, it was possible to follow
up the formation of such neuronal multisensory connections
along with piano training in early pianists. Non-musicians, who
had never played an instrument before, were trained on a computer piano twice a week over a period of five weeks. They listened to short piano melodies of a three-second duration played
in a five-tone range, and were then required, after a brief pause,
to replay the melodies with their right hand as accurately as possible. After 20 minutes of training, first signs of increased neuronal coupling between auditory and motor brain regions were
observable. After five weeks, listening to piano tunes produced
additional activity in the central and left sensorimotor regions.
In turn, playing on a mute (soundless) keyboard produced additional activity in the auditory regions of both temporal lobes.3
This experiment impressively demonstrates how dynamically
brain adaptations accompany these multi-sensorimotor
learning processes.
Activation of motor co-representations can occur in trained
pianists not only by listening to piano tunes, but also by
observing a pianists finger movements while watching a video.
The brain mechanisms of such learning through observation
have been clarified in recent years. When monkeys observed the
actions of co-species, for example grasping peanuts, exactly the
same brain areas were active as if the observing monkeys were
performing the action themselves. Additionally, a region in the
parietal lobe of the observed monkeys was activated, which is
believed to represent the knowledge that it is not me who is
performing the action. Quite appropriately, this neuronal network was termed a mirror neuron network. When trained
pianists observe video sequences of a moving hand at the piano,
the motor hand area in the primary motor cortex, secondary
auditory cortices in the temporal lobe and the cerebellum are
activated, thus impressively demonstrating such a mirrorsystem in humans.4 As a consequence for musical practice, it
follows that careful demonstration with the instrument may
enhance learning.
Practising through listening and/or observation can be considered as special cases of mental training. Narrowly defined,
mental training is understood as the vivid imagination of movement sequences without physically performing them. As with
observation of actions, principally the same brain regions are
Clinical Medicine Vol 8 No 4 August 2008
Royal College of Physicians, 2008. All rights reserved.

active as if the imagined action is performed; that is, the primary

motor cortex, the supplementary motor cortex and the cerebellum.5 In a study investigating mental training of finger movement sequences of different complexities, brain activation
increased along with the degree of difficulty of the imagined
motor task. Furthermore, when continuing mental practice over
a period of several days, the involved brain regions showed
plastic adaptations. Although these adaptations are less dramatic than if the motor tasks were practiced physically, mental
training produced a clear improvement in task performance as
assessed in finger-tapping tests.

Plasticity of sensorimotor systems: musicians

brains are different
During the past decade, brain imaging has provided important
insights into the enormous capacity of the human brain to
adapt to complex demands. Brain plasticity is best observed in
complex tasks with high behavioural relevance for the individual such that they cause strong emotional and motivational

Increased brain activation in professional pianists compared to

age-matched non-musicians while:

Listening to
piano tunes

Speaking in sounds
Playing on
a soundless
keyboard
with right
hand

The interface

Fig 1. Increased brain activation in professional pianists

compared to age-matched non-musicians. Additional brain
activity (yellow/red zones) in skilled pianists compared to nonpianists when listening to piano tunes without moving their
fingers (upper row), or when moving their fingers on a mute
keyboard (lower row). When listening, the primary hand area of
the pre-central area is active demonstrating an unconscious corepresentation of heard tunes as movement patterns.
Furthermore, an auditory association area between the temporal
and the parietal lobe is lighting up (blue arrows). This region is
similarly active when the pianists are moving their fingers on the
mute keyboard. Therefore, this region seems to be an auditorymotor interface, translating the sounds into fingerings and vice
versa. Moving the fingers on a mute keyboard (lower row, left)
additionally produces an activity in Brocas area. Therefore, this
area is not confined to language functions as has been
traditionally believed, but also contributes in a more general
manner to complex movement patterns which code symbolic
communication. It impressively demonstrates that music speaks
to our souls. Reprinted with permission from Elsevier.2

411

Eckart Altenmller

activation. Plastic changes are more pronounced in situations

where the task or activity is intense and the earlier in life it has
been developed.
Comparison of the brain anatomy of skilled musicians with
that of non-musicians shows that prolonged instrumental practice leads to an enlargement of the hand area in the motor cortex
and to an increase in grey matter density corresponding to more
and/or larger neurons in the respective area.6,7 These adaptations appear to be particularly prominent in all instrumentalists
who have started to play prior to the age of 10 and correlate positively with cumulative practice time. Similar effects of specialisation have been found with respect to the size of the corpus
callosum. Professional pianists and violinists tend to have a
larger anterior (front) portion of this structure, especially those
who have started prior to the age of seven.8 Since this part of the
corpus callosum contains fibres from the motor and supplementary motor areas, it seems plausible to assume that the high
demands on coordination between the two hands, and the rapid
exchange of information may either stimulate the nerve fibre
growth the myelination of nerve fibres that determines the
velocity of nerve conduction or prevent the physiological loss
of nerve tissue during aging.
It is not only motor areas, however, that are subject to
anatomical adaptation. By means of magnetoencephalography
(MEG), the number of nerve cells involved in the processing of
auditory or somato-sensory stimuli can be monitored. Using
this technique, professional violinists have been shown to
possess enlarged sensory areas corresponding to the index
through to the small (second to fifth) fingers of the left hand
even though their left thumb representation is no different from
that of non-musicians.9 Again, these effects were most pro-

nounced in violinists who started their instrumental training

prior to the age of 10. In summary, when training starts at an
early age (before about seven years), these plastic adaptations of
the nervous system affect brain anatomy by enlarging the brain
structures that are involved in different types of musical skills.
When training starts later, it modifies brain organisation by
re-wiring neuronal webs and involving adjacent nerve cells to
contribute to the required tasks. These changes result in
enlarged cortical representations of, for example, specific fingers
or sounds within existing brain structures.

Apollos curse: musicians dystonia

There is a dark side to the increasing specialisation and prolonged training of musicians, namely loss of control and degradation of skilled hand movements, a disorder referred to as
musicians cramp or focal dystonia (Fig 2). The first historical
record, from 1830, appears in the diaries of the ambitious
pianist and composer Robert Schumann.10 As was probably the
case for Schumann, prolonged practice and pain syndromes due
to overuse can precipitate dystonia, which develops in about 1%
of professional musicians and in many cases ends their career.11
Neuroimaging studies point to dysfunctional (or maladaptive)
neuroplasticity as one of the relevant pathomechanisms.

D5
D2D4
D1

Affected digits
D3D5
D1

D2

Non-dystonic hand

Fig 2. Symptoms of dysfunctional plasticity: typical patterns of

dystonic postures in a pianist, a violinist, a flutist and a
trombone player. Most frequently, involuntary curling of fingers
and compensatory extension of adjacent fingers is observed. In
wind instrumentalist, dystonia involves sensorimotor control of
the embouchure. Typically in dystonia, no pain or sensory
symptoms are reported. Dystonia may afflict almost all groups of
instrumentalists but is more frequently seen in the right hand of
guitarists and pianists and in the left hand of violinists. Reprinted
with permission from Elsevier.12

412

Fig 3. Neuronal correlates of dysfunctional plasticity. Fusion of

the somatosensory representation of single digits of the hand in
a musician suffering from focal dystonia. The best fitting dipoles
explaining the evoked magnetic fields following sensory
stimulation of single fingers are shown projected on the
individuals magnetic resonance image. Whereas for the nonaffected hand the typical homuncular organisation (see inset)
reveals a distance of about 2.5 cm between the sources for the
thumb and the little finger (white square and brown circle on the
left), the somatosensory representations of the fingers on the
dystonic side are blurred, resulting from a fusion of the neural
networks which process incoming sensory stimuli from different
fingers (red circles). Modified with permission from References 1
and 13.

Clinical Medicine Vol 8 No 4 August 2008

Royal College of Physicians, 2008. All rights reserved.

Neurology of musical performance

Support for this theory comes from a functional brain imaging

study performed in musicians with focal dystonia. Compared to
healthy musicians, musicians with dystonia showed a fusion of
the digital representations in the somatosensory cortex, reflected
in the decreased distance between the representation of the index
finger and the little finger when compared to healthy control
musicians.13 An example of a dystonic musician is shown in Fig 3.
Such a fusion and blurring of receptive fields of the digits may
well result in a loss of control, since skilled motor actions are
necessarily bound to intact somatosensory feedback input.
Considering (a) the historical advent of the disorder in the 19th
century with rapidly increasing technical demands imposed on
musicians, (b) the epidemiological data with rapid and repetitive
finger movements as a risk factor, and (c) the neurobiological
findings of the blurring of hand representations, it would be
tempting to state that focal dystonia finally marks the natural
limits of a process of refinement of manual dexterity over a million years. However, hereditary factors with a certain predisposition to develop this condition may also play a role.14
In summary, musical performance is an excellent model to
study the effects of neuroplasticity in the auditory and the
sensorimotor domains. It seems to be one of the most powerful
stimuli to drive plastic changes in the CNS. Investigating professional musicians might differentiate the contributions of experience or training from those of genetic predisposition. Studying
focal dystonia finally, can illustrate the effects of dysfunctional
plasticity due to overuse. An important question arises from the
investigations presented here. One has to bear in mind that
essential to music is to elicit strong emotional reactions. Shivers
down the spine, tears in the eyes, a lump in the throat while
listening to music are accompanied by the activation of a brain
network which is involved in reward, emotion and motivation.
Further research is required to show whether activity in these
areas is mediating the powerful effects on neural plasticity.

2 Bangert M, Peschel T, Rotte M et al. Shared networks for auditory and

motor processing in professional pianists: Evidence from fMRI
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3 Bangert M, Altenmller E. Mapping perception to action in piano
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4 Haslinger B, Erhard P, Altenmller E et al. Transmodal sensorimotor
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10 Altenmller E. The end of the song: Robert Schumanns focal
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11 Jabusch HC, Altenmller E. Epidemiology, phenomenology and
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References
1 Mnte TF, Altenmller E, Jncke L. The musicians brain as a model
of neuroplasticity. Nat Neurosci Rev 2002;3:4738.

Clinical Medicine Vol 8 No 4 August 2008

Royal College of Physicians, 2008. All rights reserved.

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