144

Ann. N.Y. Acad. Sci. 999: 144-151 (2003). 2003 New York Academy of Sciences.
doi: 10.1196/annals.1284.016
Neuropsychological Studies of Musical Timbre
SEVERINE SAMSON
University of Lille 3, URECA, and Epilepsy Unit, Salptriere Hospital, Paris, France
ABSTRACT: Musical timbre is a multidimensional property of sound that allows
one to distinguish musical instruments. In this paper, studies that explore the
cerebral substrate underlying the processing of musical timbre are discussed.
Perceptual asymmetries measured in normal participants, deficits of musical
timbre perception obtained in brain-damaged patients, as well as results
obtained with various neuroimaging methods are reviewed. The findings
obtained in all of these studies generally support the predominant involvement
of right temporal lobe areas, and more specifically of its anterior part, in pro-
cessing spectral and temporal envelopes of musical timbre. However, contro-
versies still exist about the contribution of the left temporal lobe in timbre
perception. The necessity of comparing data obtained with different perceptual
paradigms (same-different discrimination and similarity judgment) and vari-
ous types of stimuli (single tones and melodies) was emphasized by reporting
lesion studies carried out in patients with unilateral temporal lobe lesions. The
few neuroimaging studies published in this domain provided additional and
complementary findings. Unlike lesion studies that allow us to infer the cere-
bral structures that are essential for timbre perception, the latter investigations
implicate a more distributed neural network in timbre processing that extends
along the superior temporal gyrus to include not only anterior but also poste-
rior temporal regions and possibly frontal areas as well.
KEYWORDS: musical timbre; perception; brain
INTRODUCTION
A growing interest in the psychology oI music and in neuropsychology has been
devoted to the study oI musical timbre, a multidimensional property oI sound that
allows a person to distinguish musical instruments when pitch, loudness, and per-
ceived duration remain identical.
1
Unlike these latter acoustical attributes that
depend on a single physical dimension, timbre is deIined as a combination oI various
acoustical parameters involving stationary (e.g., steady-state spectral energy) and
dynamic (e.g., attack and decay times) Ieatures oI tone.
26
The covariation oI these multiple cues leads timbre to be more distinctive than
pitch or duration. Indeed, it has been shown that nonmusicians are more sensitive to
changes in timbre than to changes in pitch in a sound categorization task.
7
A number
oI studies also showed that diIIerences in timbre are very eIIective initiators oI
Address Ior correspondence: Dr. Severine Samson, Universite de Lille 3, URECA, UFR de
Psychologie, BP 149, 59653 Villeneuve d`Ascq cedex, France. Voice: 33 3 20 41 64 43; Iax:
33 3 20 41 63 24.
samsonuniv-lille3.Ir
145 SAMSON: MUSICAL TIMBRE
stream segregation,
8,9
being oIten more powerIul than segregation based on pitch
diIIerences.
6,10,11
Given the importance oI timbre in perception and organization oI
the auditory scene, it seems relevant to improve our knowledge oI the neural network
underlying musical timbre.
To explore the cerebral structures involved in timbre, laterality and lesion studies
as well as brain imaging investigations were carried out. In the present paper, per-
ceptual asymmetries in normal participants and deIicits oI musical timbre perception
obtained in brain-damaged patients as well as results obtained with various neuro-
imaging methods are reviewed.
LATERALITY STUDIES
Cerebral lateralization oI musical timbre was investigated by interpreting percep-
tual asymmetries tested with dichotic or monaural stimulation. Generally, ear advan-
tages Ior nonverbal stimuli appear to be less robust than those Ior verbal stimuli.
Indeed, some studies Iailed to detect ear asymmetries Ior timbre processing.
1215
However, leIt-ear advantage (LEA), indicative oI a right-hemisphere superiority,
was reported in several studies using natural instruments,
1618
synthetic sounds,
19,20
and realistic or unrealistic (played backward) musical instruments.
17
All oI these
laterality studies strongly support a right-hemisphere priority Ior timbre perception.
LESION STUDIES
The Iirst study that examined the eIIect oI a unilateral temporal lobe lesion in
timbre discrimination was carried out by Milner
21
in 1962. Using the Seashore mu-
sical battery,
22
she showed that compared to patients with leIt temporal lobectomy,
patients who had undergone right temporal lobectomy Ior the relieI oI intractable
epilepsy demonstrated a signiIicant deIicit in discriminating spectral inIormation oI
harmonic tones, suggesting that processing oI spectral inIormation involved in tim-
bre relies on the integrity oI the right superior temporal lobe. Convergent evidence
was reported in several subsequent studies on timbre perception in surgical cases or
vascular stroke patients.
2326
However, a more recent study Iailed to replicate
Milner`s Iinding.
27
No impairment on the Seashore musical battery was obtained in
patients tested beIore or aIter unilateral temporal lobe surgery Ior epilepsy. The
authors attributed the divergent results to diIIerences between studies in the extent
oI temporal lobe resection along the Sylvian Iissure. Whereas in Milner`s study a
large portion oI the superior temporal gyrus was removed, their resection either
spared or was limited to its most anterior part, indicating that the auditory areas im-
plicated in timbre processing were still Iunctional in this latter study.
In only one study, to our knowledge, was a leIt-hemisphere advantage Ior timbre
processing Iound in lesion studies.
24
The investigators interpret the leIt-hemisphere
superiority obtained in patients with commissurotomy during identiIication oI
musical instruments as evidence oI the leIt-hemispheric semantic processing
involved in a verbal identiIication task. However, this eIIect may also be attributable,
at least in part, to the role oI the leIt hemisphere in processing timing cues reported
in several studies.
2834
Since temporal inIormation given by the initial part oI the
146 ANNALS NEW YORK ACADEMY OF SCIENCES
sound seems to be important Ior the recognition oI musical instruments,
3537
recog-
nition deIicits oI musical instruments may be aIIected by deIicits in perception oI
temporal inIormation involved in timbre.
DISCRIMINATION OF SPECTRAL AND TEMPORAL ENVELOPES
OF TIMBRE
To speciIy the role oI each temporal lobe in perceiving the spectral and temporal
cues involved in musical timbre, Zatorre and I tested patients with unilateral tempo-
ral lobe lesions involving the anterior part oI the lateral temporal neocortex as well
as various extent oI the medial temporal lobe structures in a discrimination task.
38
Tones with varying spectral or time envelopes were created by changing the number
oI harmonics or the rise time duration, respectively. HalI the stimuli consisted oI
complex tones with 3, 4, or 5 harmonics with a constant temporal envelope (1-ms
rise time). The other halI had rise times oI 1, 50, or 190 ms with a constant spectral
content (4 harmonic tones). Tones were presented in pairs, and the subject had to
respond whether the two tones were the same or diIIerent. The results showed that
right but not leIt temporal lobectomy impaired the discrimination oI tones diIIering
in their spectral envelope or their temporal envelope, suggesting that the right tem-
poral lobe is essential Ior processing the temporal as well as the spectral inIormation
involved in musical timbre.
Although right temporal lobe superiority in processing spectral inIormation
conIirmed previously reported data, the predominant role oI the right temporal lobe
in processing time-related cues was a new Iinding. It seems to contradict previous
results showing that leIt hemisphere structures may be predominantly involved in
temporal processing.
30,32,3941
However, the temporal Ieatures manipulated in these
other studies involve rapid temporal coding presented sequentially. In our investiga-
tion, the discrimination oI temporal inIormation is generally based on slower tempo-
ral changes that holistically modiIied tone perception. From our Iindings, we can
conclude that timbre discrimination oI spectral energy and oI temporal variations
depends on the integrity oI auditory areas Irom the right temporal neocortex.
DISSIMILARITY 1UDGMENT OF MUSICAL TIMBRE
Recently, we have used another paradigm involving similarity judgments to study
musical timbre perception in single tones as well as in melodies.
42
Similarity judg-
ment is based on the extraction oI particular Ieatures that play an important role in
recognition by requiring higher order perception than discrimination or detection oI
diIIerences.
43
The participants included in this study were also tested with a conven-
tional 'same-diIIerent task,
38
thereby allowing the comparison oI two perceptual
procedures. For this purpose, synthetic sounds were created by selecting three levels
oI spectral changes (1, 4, and 8 harmonics) and three levels oI temporal changes (1,
100, and 190-ms rise time duration). All oI these tones were diIIicult to label, there-
Iore precluding the use oI verbal strategies. Pairs oI diIIerent stimuli were presented
to patients with unilateral temporal lobe excisions and normal control participants Ior
judgment oI their similarity on a rating scale. This task, already validated in normal
147 SAMSON: MUSICAL TIMBRE
participants,
44
provides an opportunity to examine the dependence oI timbre on both
spectral and temporal envelopes oI tones. The method used to analyze the data was
based on a multidimensional scaling (MDS) technique that allows us to represent the
dissimilarities between a set oI physically diIIerent stimuli as distances between
points in an n-dimensional space.
4547
MDS reveals the extent to which physical
distances between multidimensional acoustic inIormation are actually perceived by
a subject, instead oI investigating only the ability to distinguish same Irom diIIerent
pairs oI tones.
In a previous study,
44
we showed that normal participants could recover in their
perceptual judgments the unambiguous acoustic dimensions that were physically
inherent to the stimuli. More speciIically, normal listeners could perceive the num-
ber oI harmonics and rise time duration in a gradual and ordered way and to use this
inIormation to make perceptual judgments oI musical timbre in single tones as well
as in melodies. By contrast, perceptual ratings obtained by patients with right temporal
lobe lesions revealed a disturbed perceptual space in single tones and melodic condi-
tions. The most distorted results were obtained with single tones, in which the temporal
parameter was less prominent. Tones were grouped according to their spectral content,
but the results did not reIlect a coherent underlying perceptual dimension. The results in
patients with leIt temporal lobe lesions suggest that they could use the spectral and the
temporal envelopes oI tones independently in making perceptual judgments oI single
tones. In the melodic condition, their results were signiIicantly diIIerent Irom those oI
normal participants, suggesting that leIt temporal lesions do aIIect subtle aspects oI
timbre perception despite these patients` preserved ability to make discrimination
judgments using traditional paradigms. In general, data Irom both patient groups
showed that extraction oI temporal cues was easier in the melodic than in the single tone
condition, suggesting that the diIIerent durations and Irequencies heard within a musical
context Iacilitate timbre perception. The Iindings oI the present study replicate and
extend previous results showing that timbre perception mainly depends on the integrity
oI right neocortical structures. However, the contribution oI leIt temporal regions is also
apparent, indicating that multidimensional techniques are sensitive to more subtle
perceptual disturbances that may not be revealed by 'same-diIIerent discrimination
paradigms.
38
Lesion studies converge to ascribe a crucial role to the superior right temporal
lobe regions in timbre perception. This right hemisphere bias seems to result Irom
the dysIunction oI the anterior temporal lobe structures rather than Irom the dysIunc-
tion oI posterior temporal lobe areas, because these later regions are generally spared
in surgical excision oI the temporal lobe. The question regarding the possible
involvement oI leIt hemisphere structures in timbre processing remains open.
BRAIN IMAGING STUDIES
The cerebral substrate associated with timbre processing has also been examined
using various neuroimaging techniques. However, the goals oI these studies were
very diIIerent, making comparison between studies diIIicult. In electroencephal-
ography studies, cortical auditory evoked potentials in response to diIIerent timbres
were explored. The results showed that larger electrical activities were recorded over
the right than over the leIt hemisphere at temporal electrodes
4850
and, to a lesser
148 ANNALS NEW YORK ACADEMY OF SCIENCES
degree, at Irontal electrodes,
48
suggesting the existence oI a predominant right hemi-
sphere neural network responding to timbre changes.
In another study
5
using magnetoencephalography, the eIIect oI musical training
on auditory cortical representations in response to tones oI diIIerent instruments
(trumpet and violin) was examined. In this study, neuromagnetic measures were
recorded in skilled musicians (trumpeters and violinists) when listening to sine tones
or musical tones played on their instrument oI training or on a diIIerent instrument.
These investigators recorded larger dipole moments in both right and leIt temporal
regions Ior tones played on a musical instrument than Ior sine tones, this eIIect being
more salient Ior the instrument oI training. Although this Iinding does not speciIy
the electrophysiologic responses speciIic to musical timbre, it suggests that a certain
degree oI neural plasticity associated with musical timbre has taken place over the
right and the leIt hemisphere in skilled musicians.
Two positron emission tomography studies were also carried out in this domain.
The Iirst one demonstrated bilateral parietotemporal activations with right greater
than leIt Irontotemporal asymmetries
52
using the timbre subtest oI Seashore`s bat-
tery.
22
More recently, Platel et al.
53
identiIied activations in the superior and middle
Irontal gyrus within the right hemisphere during the detection oI timbral changes.
However, these Ioci oI activity are diIIicult to attribute purely to timbre processing,
because variations in pitch and rhythm were also present, requiring selective atten-
tion to the timbre dimension. Nevertheless, converging evidence has been obtained
with Iunctional magnetic resonance imaging (IMRI). One oI these studies was
designed to examine auditory imagery oI musical timbre. To veriIy iI timbre imagery
shows the right-sided bias generally reported in timbre perception, Halpern and her
collaborators
54
studied the neural substrate underlying timbre perception and mental
imagery oI musical instruments. The subject`s task consisted oI making similarity
judgments oI musical instruments, either played or imagined. Consistent with lesion
studies, the perception task activated several areas in the anterosuperior temporal
gyrus with a right-sided asymmetry. When the judgments concerned not only heard
timbres but also imagined timbres, posterior temporal areas were activated, relative
to the visual imagery control condition, again with a slight right hemisphere predom-
inance. The imagery task also activated leIt Irontal areas that were attributed by the
authors to the working memory component oI maintaining the auditory image cued
by the names oI the musical instruments. This pattern oI results showing that timbre
similarity judgments oI single tones appear to be mediated largely by the right tem-
poral lobe structures conIirms and extends previous results obtained in patients
during a perceptual task.
42
The other IMRI study was designed to speciIy the char-
acteristics oI cerebral responses associated with diIIerences in melodic stimuli based
on timbre.
55
The two timbres used in this study were taken Irom extreme positions
oI a psychological space
56
and diIIered in terms oI various acoustic cues (e.g., attack
time, spectral centroid, and spectral Ilux). Throughout the experiment, listeners were
exposed to blocks oI diIIerent melodies played in either one timbre or the other.
Because both experimental conditions involved listening to timbres, the neural net-
work shared by the two conditions was cancelled out by the subtractive method, and
the pattern oI activation characterizing timbre perception in general could not be
visualized. However, the investigators identiIied bilateral activations speciIic to one
oI the two timbres (Ior instance, timbre A), as compared to the other (timbre B), in
the superior temporal gyrus (dorsomedial surIace) and the superior temporal sulcus.
149 SAMSON: MUSICAL TIMBRE
Conversely, when activations speciIic to timbre B compared to timbre A were exam-
ined, the authors Iound no signiIicant results, suggesting that each area activated by
timbre B was also activated by timbre A. This surprising pattern oI activations was
interpreted by the authors as resulting Irom the rapid temporal changes in spectral
energy distribution that characterized timbre A. However, we also suggest that the
melodic context in which timbre is presented may have inIluenced timbre percep-
tion, as we already pointed out,
42,44
by obscuring or enhancing the importance oI
certain parameters that may recruit the leIt temporal lobe areas in addition to those
on the right.
CONCLUSIONS
The Iindings obtained in lesion and brain imaging studies generally support the
involvement oI the right auditory temporal areas, and more speciIically oI its anterior
part, in processing spectral and temporal envelopes oI musical timbre. However,
controversies still exist about the possible contribution oI leIt temporal lobe struc-
tures in timbre perception. Although we must remain cautious in interpreting these
data, the results oI lesion studies emphasize the necessity oI comparing Iindings
obtained with diIIerent perceptual paradigms and various types oI stimuli. We
demonstrated the relevance oI evaluating perceptual ratings oI similarity judgments
to inIer the physical distances between stimuli that are actually perceived by the
patient, instead oI assessing only the ability to diIIerentiate same Irom diIIerent pairs
oI stimuli in a discrimination task. Similarly, we showed that similarity judgments
oI musical timbre can be diIIerently aIIected in single tones compared to melodies,
the relative weighting oI the various acoustical Ieatures being diIIerent in these two
conditions. Finally, cerebral activations obtained in neuroimaging and deIicits
resulting Irom brain damage do not provide exactly the same inIormation. As report-
ed in several brain imaging studies, leIt temporal lobe regions might well be involved
in timbre processing, but it does not mean that these structures are essential Ior this
cognitive ability. Unlike lesion studies that permit inIerences about the indispensable
structures Ior a speciIic cognitive processing, imaging studies in normal participants
allow identiIication oI cerebral structures normally involved in this Iunction, which
include not only anterior but also posterosuperior temporal lobe areas.
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