I MEDICINE, MUSIC AND THE MIND

Do musicians have different brains?

Lauren Stewart

Lauren Stewart ABSTRACT – The search for anatomical correlates high resolution images of the living brain. Statistical
BA MSc PhD, of special skills dates from the end of the 19th approaches now allow precise quantification of dif-
Lecturer, century, when post-mortem brains of gifted indi- ferent aspects of brain structure, making it possible
Department of viduals, including musicians, were examined for to determine whether groups of individuals with
Psychology,
clues as to origins of their prized abilities. special skills exhibit anatomical specialisations.
Goldsmiths,
Modern neuroimaging techniques provide the These technical advances have been accompanied
University of
London
chance to interrogate the brains of living musi- by a revolution in the understanding of the brain’s
cians. Structural and functional specialisations capacity to change in response to experience. Only a
Clin Med have been demonstrated across several sensory, few decades ago, the mature brain was considered to
2008;8:304–8 motor and higher order association areas. These be hard-wired and immutable but animal and
specialisations are often instrument- or effector- human studies have now demonstrated considerable
specific and correlate with aspects of the training capacity for reorganisation following deafferentation
history supporting the view that they are the or stroke; environmental enrichment or learning.
result, rather than the cause, of skill acquisition. The reconceptualisation of the brain as a malleable
Musicians constitute a model, par excellence, for system forces us to consider the direction of causality
studying the role of experience in sculpting brain in cases where anatomical features are linked to the
processes. A key challenge for the future will be possession of musical, artistic or intellectual gifts.
to develop theoretical frameworks within which While these features would once have been consid-
musicians and other occupationally specialised ered to be innately determined, evidence of the
groups can be studied in order to investigate the brain’s plasticity now invites an alternative view. The
nature, scope and limits of neuroplasticity. following review summarises the structural and
functional specialisations exhibited by musicians,
KEY WORDS: adaptations, learning, musical, skill and presents evidence to suggest that they are the
result, rather than the cause, of skill acquisition.

In a museum in Salzburg hangs a sketch of two ears,

Altered motor and somatosensory maps
dated from the 16th century. The one on the left
in musicians
belongs to one of the most eminent musicians in
recorded history, the one on the right is that of an A professional keyboard player can produce up to
‘ordinary’ individual. The exhibition notes do not 1,800 notes per minute, with a precision in space and
reveal whether scholars of the time made anything of time that is unsurpassed in any other sphere of
the larger lobe or more gradual curvature of the ear human behaviour.2 This expertise in fine finger con-
on the left but it is hard not to see the sketch as an trol has a correlate in the brain. A morphometric
attempt to shed light on the basis of Mozart’s prodi- study revealed that the intrasculcal length of the pre-
gious abilities. The role of the brain in mental central gyrus (ILPG), a marker for the cortical motor
processes was established in the 19th century, after hand area, is longer in keyboard players relative to
which time the origins of ‘gifts’, including music, non-musician controls.3 Although it is possible to
were sought in the cortex. Auerbach examined the propose that individuals born with a longer ILPG
post mortem brains of several notable musicians and will have greater aptitude for playing the keyboard
noted peculiarities in temporal and parietal areas compared with those with a shorter ILPG; the asso-
that were hypothesised to account for the superior ciation between ILPG and the age at which training
musical skills seen in these individuals.1 In the commenced suggests that anatomical differences in
absence of rigorous statistical methods, however, it motor cortex are the result, not the cause of learning.
was impossible to determine whether these varia- By exploiting the differences in motor demands
tions were anything other than individual differences required for different instruments, it is possible to
in brain configuration. The development of modern further explore the relationship between anatomy
techniques such as magnetic resonance imaging (MRI) and expertise. In string players, for instance, there is
has made it possible to obtain three-dimensional, an asymmetry in the requirements for fine finger

304 Clinical Medicine Vol 8 No 3 June 2008

© Royal College of Physicians, 2008. All rights reserved.
 Do musicians have different brains?

control between the two hands, with the left hand (right motor authors point out that the right hand is often more involved in
cortex) performing the fast finger movements and the right fast finger movements than the left, which often has an accom-
hand (left motor cortex) performing bowing. In pianists, there panying role. Instrument-specific differences in anatomy would
is a more equal division of labour, with fine finger control seem to provide further support for the view that anatomical
required of both hands. A recent study compared motor cortex specialisations in musicians are use-dependent. The alternative
anatomy between keyboard players, string players and non- view, that innately specified differences in cortical anatomy
musicians.4 The inverted omega sign or ‘hand knob’ of the pre- determine instrument choice, seems unlikely given findings that
central gyrus is a gross-anatomical feature that is associated with emphasise the significant role of environmental factors in this
the representation of finger movements. Its prominence can decision.5
range from being barely present, to clearly visible, sometimes to The expanded representations seen in the motor system in
the extent that a double omega can be discerned. Since increased musicians are paralleled by changes in the somatosensory
cortical volume results in enhanced gyral folding, the extent of system. Stimulation of the digits of the left (fingering) hand of
prominence can be used as an indirect index of the volume of string players has been shown to produce an enhanced
this motor hand area. Based on the results of raters who scruti- somatosensory evoked response that correlates with the age at
nised structural MRI images from keyboard players, string which training commenced.6 The evoked response from the
players and non-musicians, musicians were found to have a right (non-fingering) hand was no different between string
more prominent omega sign (OS) configuration than non- players and non-musicians, suggesting that the expanded cor-
musicians. While string players showed greater OS prominence tical representation results from the increased afferent input
in the right hemisphere, consistent with the fine finger control received by the fingering hand. Evidence pertaining to the likely
required of the left hand, keyboard players showed greater functional significance of the enhanced somatosensory
prominence in the left hemisphere. While a bilateral promi- responses comes from a study in which pianists were found to
nence may have been expected in the keyboard group, the have improved tactile sensitivity of both hands compared with
non-musician controls, correlating with duration of daily prac-
tice.7 The effector specificity and the relationship between the
size of the effect and aspects of the training history in these two
studies supports a use-dependent view of these specialisa-
M(h) SP tions.6,7 Both these paradigms would lend themselves to longi-
S(h) tudinal designs, whereby evoked responses and tactile sensitivity
could be measured during the course of learning. Emergent
behavioural and neurophysiological differences learning would
PT be irrefutable evidence for the causal role of learning and such
HG an approach would also shed light on the time course of these
changes.
IF IT

Increased interhemispheric processing in

musicians
Fine finger control is of critical importance for skilled musical
performance, but of equal significance is the ability to coordi-
nate sensorimotor processing across the effectors – not only
hands, but sometimes feet, lips and respiratory muscles.
Coordination between the hands is the simplest case of sensori-
motor coordination, requiring extensive neural transmission
between the hemispheres. Evidence from morphometric studies
suggests that interhemispheric connections are enhanced in
CC
musicians. Professional musicians who commenced their
training before age seven were found to have a larger anterior
portion of the corpus callosum compared to non-musician con-
trols (see Lee et al for an interaction with gender that remains to
be explored).8,9 While suggestive of enhanced interhemispheric
communication, structural differences alone say nothing about
Fig 1. A schematic representation showing regions of the functional role of these anatomical differences. A transcra-
specialisation in the brains of musicians. CC = corpus callosum; nial magnetic stimulation (TMS) study investigated the inter-
HG = Heschl’s gyrus (site of primary auditory cortex); IF = inferior
frontal cortex; IT = inferior temporal cortex; M(h) = hand area of hemispheric communication in musicians and non-musi-
motor cortex; PT = planum temporale; S(h) = hand area of cians.10 Transcranial magnetic stimulation involves the delivery
somatosensory cortex; SP = superior parietal cortex. of a magnetic pulse to the brain via a coil placed against the

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© Royal College of Physicians, 2008. All rights reserved.
Lauren Stewart

scalp. The induced electric field depolarises neurons underlying that the enhancement emerges from a striving to produce or
the coil, in the surface of the cortex. When given over the hand modulate the timbral qualities of one’s instrument.
area of the primary motor cortex, this elicits muscle contraction The obligatory coupling between sound and action in musical
in the contralateral hand that can be measured via surface elec- performance is well known to musicians. Singers who need to
trodes applied to the first dorsal interosseus muscle. When a rest their voices before a performance are instructed not to listen
‘conditioning’ pulse is given over the ipsilateral motor hand area to music, in order to avoid straining the voice through auto-
before the ‘test’ pulse is given, the muscle contraction is of lower matic subvocalisation (Collyer, personal communication, 2007).
amplitude, owing to the activation of inhibitory circuits Two recent functional (f)MRI studies compared activation
between the motor hand areas of each hemisphere. This inter- when musicians listened to a piece of music without playing it
hemispheric inhibition is known to have an important role in versus when they played a piece of music without auditory feed-
motor control, for instance, by preventing mirroring move- back.18,19 Both studies showed areas of overlap between the two
ments and allowing the effectors to operate independently of conditions, including premotor cortex, supplementary motor
one another. Musicians, however, were shown to have reduced area and planum temporale. A magnetoencephalography
interhemispheric inhibition, since the conditioning pulse was (MEG) study showed that purely listening to music which was
less effective in reducing the amplitude of muscle contraction in within the listeners repertoire resulted in a response from pri-
response to the test pulse compared with non-musician con- mary motor cortex.20 Moreover, a dissociation in evoked
trols.10 At first sight this may seem counterintuitive since the response was seen between those notes which would have been
ability to independently control the two hands, is a defining fea- played by the thumb and the little finger. Similarly, when
ture of pianistic skill. However, the role of interhemispheric pianists listened to a piano piece at which they were practised,
inhibition may depend on the level of expertise. The beginning they demonstrated higher motor cortex excitability than when
pianist might experience involuntary mirroring when the two they listened to a flute piece on which they had not been
hands play together, but once independence of movement has trained.21 The demonstration that perception and action can be
been achieved, there is an obvious need for coordination in the closely coupled through musical performance opens possibili-
execution of these separate motor programs. Expert performers ties for using music to affect action, for instance, in promoting
may be able to accommodate and benefit from the effects of motor function recovery following stroke.22
reduced interhemispheric inhibition between their motor
hand areas because they have already achieved automatic and
Visiospatial processing in musicians
independent control of the two hands.
Musicians rely heavily on a sound to action link, but the
majority of them are also required to translate rapidly and con-
Auditory processing differences in musicians
tinually between visuospatial symbols and their associated
Sensorimotor processing in musicians subserves one over- actions, with a precision and fluidity that allows them to keep in
arching goal: to produce sound. Musicians are intensely attuned time with their own predetermined beat, or the conductor’s
to the sounds that they produce, employing feed-forward con- baton. The sequencing aspect of music reading appears to have
trol in order to refine and modulate the sounds produced until a correlate in left inferior frontal cortex. Two groups reported
what they hear corresponds to an internal model of the desired increased grey matter in this area23,24 and one of these groups
sound.11 Musicians have been shown to have more gray matter additionally showed that musicians, but not non-musicians,
than non-musicians in anterior medial Heschl’s gyrus (HG), activated this area while performing a non-musical sequencing
corresponding to primary auditory cortex.16 A subgroup of task.25 Musician-specific differences have also been found in
musicians who possess absolute pitch (AP), the ability to iden- superior parietal cortex and inferior temporal gyrus, which are
tify tones by name, in the absence of a reference pitch have fur- thought to reflect adaptations to different aspects of the music
ther differences in planum temporale, an auditory association reading process.24 Musical notation consists of spatial and feat-
area.12 Two studies suggest that AP possessors show an exagger- ural information, respectively conveying the ‘what’ and ‘when’
ation of the normal leftward asymmetry of this region, brought of the musical response. A longitudinal fMRI study, in which
about by a reduction in cortical volume in the right hemisphere, musically-naive adults were taught to read music and play piano
as opposed to an increased volume in the left.13,14,15 over a period of three months, revealed different functional
At a functional level, musician-specific differences have been changes, associated with the learning of each of these aspects of
found in the early and late components of the auditory evoked musical notation.26,27 When participants were required to read
response, following presentation of pure and complex tones music for pitch alone, learning-related changes were seen in
respectively.16,17 The latter component is influenced by the superior parietal cortex. When the task was to read music for
behavioural relevance of the spectral properties of the tones. rhythm alone, learning-related changes were seen in temporal
When violinists hear violin tones, they show an enhanced cortex. Although the experience of reading music is an inte-
response relative to when they hear trumpet tones, while the grated one, these findings reveal that different aspects of musical
reverse is true for trumpeters.17 On the one hand, this effect may notation are, at least initially, dealt with by different brain
have arisen from having had greater experience with the spectral regions, before these outputs are combined to produce a
features of one’s own instrument. It is also possible, however, response that is unified in space and time.

306 Clinical Medicine Vol 8 No 3 June 2008

© Royal College of Physicians, 2008. All rights reserved.
 Do musicians have different brains?

Musicians as a model special abilities. The emerging view places training and practise
centre stage, to the extent that musicians are now championed
Trained musicians provide the opportunity to study structural
as models par excellence for understanding use-dependent reor-
and functional plasticity associated with the acquisition of a
ganisation in the human brain. It is clear that we have now
number of sensory, motor and cognitive musical subskills but
moved beyond the question, ‘are musicians’ brains different?’ to
they also provide the opportunity to ask more generic questions
asking how these differences can shed new light on the relation-
concerning learning and plasticity. One such question concerns
ship between experience and brain function. A goal for the
the role of sensitive periods in learning. Does early learning have
future should be to develop theoretical frameworks within
a more profound impact on the development of ability com-
which musicians can be considered alongside other groups of
pared with later learning? Several studies have shown correla-
occupationally specialised or sensory-deprived individuals. By
tions between brain differences and age at which training com-
comparing and contrasting the effects of different parameters of
menced, but in most cases, individuals who start training earlier
experience on brain function, it will be possible to transcend the
have also been playing for a longer period of time, making it
situational constraints of any one group and elucidate general
impossible to disambiguate the role of training duration from
principles of learning and plasticity that may find application in
the stage at which the training commenced. A recent study, how-
questions of educational and clinical significance.
ever, compared early-trained (pre-seven years) and late-trained
(post-seven years) musicians on their performance of a novel
complex rhythmic sequence.28 Early-trained musicians showed Acknowledgments
enhanced performance, compared with late-trained musicians,
The author is supported by a grant from the Economic and Social
even though the two groups were matched for the overall dura-
Research Council. The author thanks Vincent Walsh and Christina Mo
tion of training. One interpretation of this is that the early for discussions relating to this article.
training may coincide more closely with relevant structural
changes occurring in the brain, compared with the training that
occurred later. This view finds support in a study using diffusion References
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308 Clinical Medicine Vol 8 No 3 June 2008

© Royal College of Physicians, 2008. All rights reserved.