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Behavioral and Cognitive Neuroscience Reviews
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The Effect of Music on Cognitive Performance: Insight From Neurobiological and Animal Studies
Nikki S. Rickard, Samia R. Toukhsati and Simone E. Field
Behav Cogn Neurosci Rev 2005 4: 235
DOI: 10.1177/1534582305285869

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10.1177/1534582305285869
BEHAVIORAL AND COGNITIVE NEUROSCIENCE REVIEWS
Rickard et al. / FACILITATORY EFFECTS OF MUSIC ON COGNITIVE PERFORMANCE
The Effect of Music on Cognitive Performance:
Insight From Neurobiological and Animal Studies
Nikki S. Rickard
Samia R. Toukhsati
Simone E. Field
School of Psychology, Psychiatry and Psychological Medicine, Monash University, Australia
The past 50 years have seen numerous claims that music expo-
sure enhances human cognitive performance. Critical evalua-
tion of studies across a variety of contexts, however, reveals
important methodological weaknesses. The current article
argues that an interdisciplinary approach is required to ad-
vance this research. A case is made for the use of appropriate ani-
mal models to avoid many confounds associated with human
music research. Although such research has validity limitations
for humans, reductionist methodology enables a more controlled
exploration of music’s elementary effects. This article also ex-
plores candidate mechanisms for this putative effect. A review of
neurobiological evidence from human and comparative animal
studies confirms that musical stimuli modify autonomic and
neurochemical arousal indices, and may also modify synaptic
plasticity. It is proposed that understanding how music affects
animals provides a valuable conjunct to human research and
may be vital in uncovering how music might be used to enhance
cognitive performance.
Key Words: cognition, emotion, animal model, mem-
ory, spatial performance, music therapy, music train-
ing, underlying mechanisms, arousal
The potential for music to influence cognitive perfor-
mance has intrigued researchers, therapists, and educa-
tors. The appeal of a “treatment” that is enjoyable, has a
very low risk of side effects, and is in most scenarios inex-
pensive is undeniable. Music has been used in an
attempt to ameliorate memory loss in demented
patients, to assist language acquisition in learning-
impaired children, and to enhance students’ academic
performance in examinations. Nevertheless, the claim
that music can have a significant impact on cognitive per-
formance lacks stability on both empirical and theoreti-
cal grounds. To date, the data in support of the claim are
sparse and, in many cases, unconvincing owing to meth-
235
Downloaded from bcn.sagepub.com at Monash University on June 12, 2013
odological weaknesses. In the few cases where compel-
ling data have been obtained, there is urgent need of
replication. There has also been extensive speculation
on the mechanisms by which music might impact cogni-
tive processes. This question has contributed to some of
the most exciting theoretical work in music psychology,
with implications for topics ranging from neurobiology
of emotions (Panksepp, 1998) to evolutionary psychol-
ogy (Wallin, Merker, & Brown, 2000). However, specific
hypotheses on the neurobiological mechanisms medi-
ating cognitive-enhancing effects of music are rare.
The current article will review a selection of work per-
taining to these issues within music neuroscience. A posi-
tion will be presented that animal research can shed
light on a number of shady impasses that currently exist,
providing a novel and innovative solution. It will be
argued that confounds inherent in much of the human
music psychology work can—for specific research ques-
tions—be circumvented by using appropriate animal
models. Furthermore, exploration of mechanisms
underlying behavioral effects can often be more direct
and controlled within the animal research paradigm.
The conclusions drawn from such data are clearly lim-
ited in their ethological validity for humans. Nonethe-
less, it will be argued that this in itself is of considerable
value: How animals respond to music tells us much
about the more “direct” effects of music on the brain and
can distinguish what might be distinctively “human”
from more fundamental effects of music that we might
Authors’ Note: We would like to dedicate this review to Emeritus Pro-
fessor Kim T. Ng. An inspirational mentor and friend; we are grateful
for the rhythm he provided to which each of us now composes.
Behavioral and Cognitive Neuroscience Reviews
Volume 4 Number 4, December 2005 235-261
DOI: 10.1177/1534582305285869
© 2005 Sage Publications
236 BEHAVIORAL AND COGNITIVE NEUROSCIENCE REVIEWS
share with other species. A comparative discussion of
how animals respond to music will provide the basis for
whether a “bottom-up” approach to this research
question is feasible.
HUMAN STUDIES
Effect of Music on Cognitive
Performance in Humans
Passive Exposure
The effect of passive exposure to music on cognitive
performance has been explored for decades. Although
deleterious effects have been observed (e.g., Fogelson,
1973; North & Hargreaves, 1999), the majority of studies
that have used music in a manner that does not compete
for attentional resources have demonstrated either no
impairment (e.g., Wolf & Weiner, 1972) or a facilitatory
effect of music on performance. Facilitatory effects have
been reported for a range of cognitive tasks, includ-
ing general intelligence tests (Cockerton, Moore, &
Norman, 1997), examination performance (Schreiber,
1988), arithmetic performance (Abikoff, Courtney,
Szeibel, & Koplewicz, 1996; Hallam, Price, & Katsarou,
2002; Miller & Schyb, 1989), and reading comprehen-
sion (Etaugh & Michals, 1975; Etaugh & Ptasnik, 1982;
Hilliard & Tolin, 1979; Stanton, 1975). For instance,
Abikoff et al. (1996) found a significant improvement
in arithmetic performance in children with attention
deficit/hyperactivity disorder during exposure to pre-
ferred music relative to silence or speech conditions.
Schreiber (1988) found that college students exposed to
rock music for the first 20 min of each class achieved sig-
nificantly higher examination grades than did students
in the silent control condition. Response time on a driv-
ing simulation computer program was also significantly
faster in the presence of both high- and low-intensity
dance music (Beh & Hirst, 1999).
Research has also demonstrated cognitive-enhancing
effects of exposure to anxiolytic (anxiety-reducing) or
“sedative” music. For example, Stainback, Stainback,
and Hallahan (1973) found that exposure to calming
background music (Air for the G String by J. S. Bach)
enhanced learning in educable mild to moderately men-
tally challenged students. Davidson and Powell (1986)
also found that exposure to “easy listening music” (char-
acterized by nonpercussive melodic sequences over
nondissonant choral structures) during a science class
significantly increased the time fifth-grade children
spent engaged in academic activities. In confirmation,
Cockerton et al. (1997) found that cognitive perfor-
mance was significantly enhanced during exposure to
background classical music (Chi Mai by Ennio
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Morricone for 5 min) in comparison to silence in under-
graduate students. More recently, Thompson, Moulin,
Hayre, and Jones (2005) found that exposure to Vivaldi’s
Four Seasons (Winter) during a category fluency task im-
proved performance of both healthy older adults and
Alzheimer’s patients. It should be noted, however, that
appropriate controls (for instance, for different types of
music) were not incorporated into many of these
studies, nor was the possible impact of music preference
considered.
Spatial performance. The most publicized, albeit highly
controversial, claim that exposure to stimulative music
facilitates cognitive performance derives from a tran-
sient enhancement of spatial-temporal reasoning
observed following exposure to a specific Mozart piano
sonata (Rauscher, Shaw, & Ky, 1993, 1995). Rauscher and
colleagues found that a Mozart sonata for two pianos
enhanced performance on an abstract/spatial reason-
ing task from the Stanford-Binet intelligence scale when
compared with silence, a relaxation tape (Rauscher et
al., 1993), a short story, minimalist music, or dance music
(Rauscher et al., 1995). This finding was replicated in a
number of studies (Rideout, 1997; Rideout, Dougherty,
& Wernert, 1998; Wilson & Brown, 1997; see review by
Hetland, 2000), and with similarly complex music by
other composers including Yanni (Rideout et al., 1998),
Bach (Ivanov & Geake, 2003), and Schubert (Nantais &
Schellenberg, 1999). The effect also appears to have
clinical significance because Alzheimer’s patients per-
formed significantly better on a visual-spatial task follow-
ing exposure to a Mozart sonata than following silence
(Johnson, Shaw, Vuong, Vuong, & Cotman, 2002) or
1930s popular tunes (Johnson, Cotman, Tasaki, & Shaw,
1998). Support for the “Mozart effect” has not, however,
been unanimous owing to equivocal replicability of the
findings. Although it can be argued that some studies
(e.g., Carstens, Huskins, & Hounshell, 1995;
McCutcheon, 2000; Newman et al., 1995; Stough,
Kerkin, Bates, & Mangan, 1994) have not adequately
replicated the original methodology to challenge the
effect (Rauscher & Shaw, 1998), fairly strict adherence to
the procedure used in the original studies has also
yielded some null effects (Steele, 2001; Steele, Bass, &
Crook, 1999; see review by Chabris, 1999).
Thompson, Schellenberg, and Husain (2001) found
that spatial performance was significantly better follow-
ing exposure to Mozart’s piano sonata in comparison to
an Albinoni adagio or silence. However, they argued,
and later demonstrated, that this difference was due to
differences in participants’ mood and arousal levels
(Schellenberg, Nakata, Hunter, & Tamoto, in press). In
support of their hypothesis, spatial-temporal perfor-
mance was found to depend on the tempo (which is asso-
 Rickard et al. / FACILITATORY EFFECTS OF MUSIC ON COGNITIVE PERFORMANCE
ciated with arousal) and mode (which is associated with
mood) of the Mozart sonata, with better performance
following a fast rather than slow version, and when the
sonata was in major rather than minor mode (Husain,
Thompson, & Schellenberg, 2002). A similar effect has
also been reported when participants enjoyed listening
to a Stephen King novel (Nantais & Schellenberg, 1999)
or the popular music band Blur (Schellenberg & Hal-
lam, in press). Finally, a similar improvement in a pro-
cessing speed task was also observed following exposure
to music by Mozart (Schellenberg, Nakata, et al., in
press). This task generalization suggests that the effect is
not limited to spatial-temporal performance and may
in fact simply be an interesting example of the well-
established arousal-performance relationship (Berlyne,
1967; Naatanen, 1973).
As previously indicated, and in opposition to this
arousal-mood hypothesis, cognitive performance (in-
cluding spatial-temporal performance) can also be en-
hanced by anxiolytic music (Cockerton et al., 1997;
Hallam et al., 2002; Stainback et al., 1973; Thompson
et al., 2005). For instance, Borling (1981) found perfor-
mance on a paper-and-pencil maze test to be signifi-
cantly better following sedative music (Debussy’s G minor
String Quartet) than stimulative music (Wagner’s Ride of
the Valkyries). Interestingly, coincident electroencephalo-
graph (EEG) measurements showed no difference in
alpha band activity following exposure to the stimula-
tive and sedative music. It is possible, therefore, that
some other attribute of the music, such as its capacity to
promote a positive mood, accounts for the cognitive-
enhancing effect of nonarousing music.
Regardless of the explanation, the finding that music—
in certain conditions, such as when it is enjoyed—can
facilitate cognitive performance is undeniably of consid-
erable psychological interest. Despite public misinter-
pretation and mythologizing about the “Mozart effect”
(Bangerter & Heath, 2004), there has never been any
empirical claim that the effect is specific to music by
Mozart. The remaining scientific controversy surround-
ing this effect tends to relate to the underlying mecha-
nism, rather than its existence (albeit of small effect
size). (The evidence for affective and arousal-related
mediation of music-induced cognitive benefits is further
explored in the section “Affective Mediation”).
Learning and memory. There has also been a keen inter-
est in the effect of music on learning and memory,
largely because of the reported benefits of music expo-
sure or therapy for clinical populations with memory
impairments (see also the section “Music Therapy”).
Empirical research on such populations has not been
extensive and is largely confined to short-term memory
processes. The available literature does, however,
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237
illustrate some evidence of an enhancing effect of expo-
sure to music on memory in children with normal cogni-
tive capacity. For example, Morton, Kershner, and Siegel
(1990) found that exposure to popular music prior to
learning facilitated memory capacity and reduced
distractability for a verbal dichotic listening task in male
children in comparison to performance following
silence. Hallam et al. (2002) found that primary school
children performed significantly better on a cued recall
memory task in the presence of background calming
music than in the presence of background music per-
ceived as unpleasant and aggressive, or silence. Recall
processes, however, appear to be generally impaired or
unaffected by music presented prior to encoding
(Hirokawa, 2004), during learning (Belsham &
Harman, 1977; Boyle & Coltheart, 1996; Crawford &
Strapp, 1994; Furnham & Allass, 1999; Sousou, 1997;
Thaut & de I’Etoile, 1993), or during retention testing
(Nittono, 1997), particularly if the music comprises a
vocal component (Salamé & Baddeley, 1989).
In contrast, music exposure appears to more consis-
tently enhance memory in cognitively impaired popula-
tions, with positive findings when the music stimulus is
presented prior to (Isern, 1960; Lord & Garner, 1993;
Prickett & Moore, 1991) and during (Shehan, 1981;
Staples, 1968) learning, and with a range of music types.
For example, Foster and Valentine (1998, 2001) found
that exposure to music, regardless of whether it was
familiar or novel to participants, facilitated autobio-
graphical recall in individuals with mild to moderate
dementia in comparison to performance following
exposure to a quiet control condition or cafeteria noise.
Exposure to “Big Band” music has also been shown to
improve autobiographical memory recall, mood, alert-
ness, and sociability in Alzheimer’s patients (Lord &
Garner, 1993; Sambandham & Schirm, 1995). Moreover,
those experiencing the severest impairments evidenced
the greatest improvements.
Children with learning disabilities have also demon-
strated improvements in memory task performance fol-
lowing music exposure. Shehan (1981) found that a
combination of music/visual or verbal/visual teaching
strategies facilitated short-term retention of a paired-
associate learning task in children with learning disabili-
ties. Similarly, Staples (1968) investigated the effects of
exposure to music on retention in 15 mild to moderately
learning-impaired children, using paired-associate tasks.
The findings showed that recall for words embedded
within an isochronous rhythm was significantly en-
hanced in comparison to the quiet control condition.
Moderating factors. Music is a multidimensional stimu-
lus, comprising a variety of musical elements (e.g.,
rhythm, melody, harmony, timbre, form), as well as
238 BEHAVIORAL AND COGNITIVE NEUROSCIENCE REVIEWS
extra-musical variables relating to an individual’s inter-
action with the piece (e.g., personality, musical prefer-
ence, musical training, mood, and familiarity). It is not
yet clear which characteristics of music are essential for
any cognitive-enhancing effects, and methodological
variations in experimental procedure can influence this
relationship. Inconsistencies in findings may, for
instance, be the result of differences in (a) the extent to
which music was presented obtrusively, (b) the time at
which music was presented relative to the task, (c) the
type of music stimulus, with familiar music tending to be
related to distraction, and (d) characteristics specific to
the listener (Baugh & Baugh, 1965). Music can therefore
be an unwieldy stimulus to subject to rigorous scientific
testing. A recurring limitation in human experimental
studies on the effect of music on cognitive performance
is the difficulty of controlling variables that are not of
central interest to that study. It is usually impossible to
exclude them from the study, because music is at risk of
losing its “musicality” if it is reduced to its component
elements (e.g., rhythm, melody). Furthermore, extra-
music factors such as familiarity and preference are
intrinsic to the human music experience, and it is unre-
alistic to expect that the influence of such variables on
each participant will be identical. Human responses to
music are highly idiosyncratic, and differences in
preference for, emotional responses to, and associations
with a music piece will inevitably differ across listeners.
One means of overcoming this limitation in human
studies is to give greater attention to the variety of music-
related and extra-musical variables that influence indi-
vidual responses to music (Hallam et al., 2002; Thaut,
1990). Accordingly, several studies have directly mea-
sured potentially moderating variables in an effort to
account for their impact. The efficacy of this strategy is
demonstrated well by studies that have measured how
personality interacts with the effect of music on perfor-
mance. Utilizing a mixed factorial design, Furnham and
Allass (1999) assessed extraversion/introversion using
Eysenck’s Personality Questionnaire, and then pre-
sented complex rock music, simple rock music, or
silence during a nonverbal learning task. Introverts
recalled more during the no-music condition than dur-
ing either music condition. In contrast, extraverts re-
called more with either type of music than during
silence. Similarly, Crawford and Strapp (1994) found
that introverts preferred to study and displayed signifi-
cantly higher recall in silence than with popular music
playing. This is consistent with Eysenck’s theory that
extraverts seek out further stimulation to perform at
their optimum levels in that their level of cortical arousal
is lower than that of introverts (Eysenck, 1976). More-
over, these studies indicate that when participants are
differentiated on the basis of personality (and indirectly,
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their basal arousal levels), inconsistencies across studies
may be reduced and the facilitative effects of music on
memory become more evident.
There have also been efforts to determine whether
certain characteristics of the music itself are responsible
for the cognitive gains caused by music treatments. In
fact, it has been suggested that research on effects of
music would benefit from a more “pharmaceutical”
approach, whereby the critical ingredients and dosage
are sought (Spintge & Droh, 1992). For instance, the
intensity (or “loudness”) at which music is presented has
been found to influence task performance. Norton
(1971) found that sixth graders performed better on ver-
bal tasks when exposed to loud music (71-90 dB),
whereas general academic and creative task perfor-
mance was better in the presence of soft music (51-70
dB). Turner, Fernandez, and Nelson (1996) found that
motor reaction time was quicker in the presence of
music played at a “comfortable” (70 dBA) volume than at
softer (60 dBA) or louder (80 dBA) volumes.
Tempo also appears to be a key component of
cognitive-enhancing music. Mayfield and Moss (1989)
found that exposure to slow music (60 bpm) impaired
performance of simple transposition and arithmetic
problems, whereas performance in the presence of fast
rock music (140 bpm) was similar to that under silence.
In contrast, errors (violations) made on a driving simula-
tion task were significantly greater in the presence of fast
(120-140 bpm) music than in the presence of slow (40-70
bpm) music (Brodsky, 2002). Nittono, Tsuda, and
Nakajima (2000) reported that cognitive performance
was significantly better when participants were exposed
to fast (158 bpm) in comparison to slow (60 bpm) classi-
cal music. However, the effects of exposure to fast (158
bpm) or slow (60 bpm) metronome tones on task perfor-
mance were undifferentiated, suggesting that tempo
interacts with other music factors, such as stimulus type
or preference. Finally, reading comprehension of high
school students was found to be better in the presence of
Vangelis’ To an Unknown Man, which is repetitive and has
a limited tonal range, than in the presence of silence or
Emerson, Lake, and Palmer’s Toccata, which is complex
in terms of both dissonance and rhythm (Kiger, 1989).
The cognitive-enhancing effect of music therefore
appears to also depend on music complexity, although
this factor is likely to interact with other factors, such as
task complexity, as well.
Music Therapy
Music therapy is the planned application of music to
improve social, emotional, physical, and/or intellectual
functioning in populations experiencing dysfunction of
some type (Bright, 1992; Snyder & Chlan, 1999). It has
been used in conjunction with conventional medicines
 Rickard et al. / FACILITATORY EFFECTS OF MUSIC ON COGNITIVE PERFORMANCE
in a variety of clinical populations, particularly in special
education (Wager, 2000), gerontology, and rehabilita-
tion programs (Aldridge, 1995), and is claimed to aid in
the treatment and management of neurological dis-
orders (Smith, 1986).
Empirical support for the utility of music therapy in
challenging cognitive impairment is, however, limited
(e.g., Aldridge, 1995), and much of the evidence to sup-
port its use is derived from anecdotal reports, case stud-
ies, or clinical observations (Brotons, Koger, & Pickett-
Cooper, 1997). For example, using a single case study
design, Baur, Uttner, Ilmberger, and Gunther (2000)
found that a female participant with severe declarative
memory deficits showed strong recognition for song
titles that had been paired with their respective melo-
dies. Similarly, using a small sample of elderly individuals
diagnosed with Alzheimer’s disease (AD), Carruth
(1997) suggested that music therapy improved face rec-
ognition ability. However, given that improvements were
observed in only 4 of the 7 participants exposed to
music, further confirmation is required. Therefore,
although such studies can provide a rich source of qual-
itative data that can complement interpretation of
quantitative findings, recommendations for the use of
music therapy based solely on such studies would be
premature.
Few rigorous experimental studies investigating the
utility of music as a therapeutic intervention have been
undertaken. Smith (1986) found that performance on
the Mini-Mental State Examination by female partici-
pants diagnosed with AD was significantly improved fol-
lowing musically cued reminiscence, verbally cued remi-
niscence, or music activity. Kumar et al. (1999) found a
reduction in stress, as indexed by blood serum levels of
arousal hormones, following 20 music therapy sessions
in male participants diagnosed with AD. However, nei-
ther study included nontherapeutic controls with which
to compare the effects of the intervention, making it dif-
ficult to exclude a Hawthorne effect (that is, where intro-
duction of nonspecific factors relating to the research is
responsible for the observed change). In addition,
music therapy research has suffered from poorly oper-
ationalized dependent variables and insufficient control
of moderating variables that relate to the population
receiving therapy (Koger, Chapin, & Brotons, 1999).
Nevertheless, there seems to be a sufficient groundswell
of positive findings in this field to justify further research
of this type (Brotons & Marti, 2003; Hanser, 1985;
Sambandham & Schirm, 1995; Savarimuthu & Bunnell,
2002; Watkins, 1997).
Music Training
A substantive body of research has accumulated on
the impact of music training and education on cognitive
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239
performance, particularly in the developing brain. The
effects of music training on academic abilities, such as lit-
eracy and numeracy, have been widely studied in chil-
dren receiving regular music lessons compared with
those who do not. The type of music training has typi-
cally been either Kodály lessons (which emphasize sing-
ing and listening accurately, use of hand signals, and
structured learning of music notation and rhythmic pat-
terns) or private instrumental lessons (chiefly key-
board). Mathematical performance was found to be
improved in 6-year-old children given Kodály music
training for 7 months (Gardiner, Fox, Knowles, & Jeffrey,
1996) and in second grade children given 6 months of
piano lessons (Graziano, Peterson, & Shaw, 1999).
Cheek and Smith (1999) also found mathematical per-
formance to be significantly better in students who had
had 2 years of private music lessons than those who had
not received music lessons. With regard to literacy, pri-
mary school children (mean age 7 years) receiving
Kodály classes 5 days a week for approximately 40 min
per session were found to perform significantly better on
reading tests than matched students not receiving
Kodály instruction (Hurwitz, Wolff, Bortnick, & Kokas,
1975). Prekindergarten children receiving regular
music lessons performed better on prereading and
writing skills compared to children who received no
music lessons (Standley & Hughes, 1997).
An increase in general intelligence (IQ) has also been
associated with music training (Costa-Giomi, 1999;
Lynn, Wilson, & Gault, 1989; Schellenberg, 2004). In
one of the most convincing studies in this field,
Schellenberg (2004) evaluated the effects of a 36-week
music program (keyboard or voice) on general IQ in 6-
year-old children. Children were randomly allocated to
groups, and control groups received either no addi-
tional lessons or drama lessons, which involved similar
nonmusic skills as the music lessons (for instance, with
students having to devote time to practice and develop
their memorization ability). The findings revealed that
children receiving either keyboard music lessons or
Kodály voice training lessons showed significantly
greater improvements in general IQ than did children
receiving drama lessons or no lessons. Children with
learning disabilities have also demonstrated improve-
ments in memory following music training. For exam-
ple, Ross (1971) found that involvement in a 6-week spe-
cialized music education program utilizing Kodály
methods significantly facilitated paired-associate learn-
ing in learning-impaired children. Keyboard instruction
has been shown to enhance spatial-temporal task perfor-
mance in children enrolled in kindergarten to third
grade (Graziano et al., 1999; Rauscher, 2002), bene-
fits from which were shown to persist for up to 2 years
(Costa-Giomi, 1999). Interestingly, the type of cognitive
240 BEHAVIORAL AND COGNITIVE NEUROSCIENCE REVIEWS
task amenable to improvement by music instruction has
been shown to vary. For example, children aged between
4 and 6 years who received 30 weeks of Kindermusik for
Young Children performed significantly better than con-
trol children in abstract reasoning, but not in pattern
analysis or vocabulary scores (Bilhartz, Bruhn, & Olson,
1999), and therefore it is advisable that researchers
adopt multiple dependent measures in studies of this
nature.
Although this avenue of research is promising, there
are major caveats that limit the conclusions that can be
drawn from the majority of music training studies. First,
many of the designs are correlational, generating spuri-
ous relationships between music training and cognitive
performance. Accordingly, better performance of
music-trained children than musically untrained coun-
terparts may be due to a third common factor, such as
higher socioeconomic status of parents, advanced
resources or extracurricular activities provided by the
school, or higher prior IQ, rather than to music training.
As with research that explores passive music exposure
and music therapy, research on the potential benefits of
music training must either exclude the contribution of
such potentially confounding factors or use an experi-
mental design in which children of similar socio-
economic status, baseline ability, and curriculum are
a l l o c a t e d to th e m u s i c an d c o n t r o l g r o u p s
(Schellenberg, 2004).
Second, many of the experimental studies have inade-
quate control conditions. That is, they provide music les-
sons to some children and compare their performance
with that of children not given music lessons (e.g., Costa-
Giomi, 1999; Hetland, 2000; Hietolahti-Ansten &
Kalliopuska, 1990). Any difference could be attributable
to a Hawthorne effect, whereby introduction of a new
teacher, an increase in teacher attention, or motivation
to work toward a performance goal is responsible for the
change rather than the music training per se. It is impor-
tant that control conditions comprise similarly engaging
and novel activities to enable exclusion of such con-
founds in interpretation (Schellenberg, 2001). The use
of appropriate controls also enables conclusions to be
drawn about the mechanism underlying any effect. For
instance, many of the skills associated with music train-
ing (e.g., disciplined practice, memory, teamwork,
pride, and so on) are likely to be shared by a suitable con-
trol activity. Should music training produce benefits not
shared by another similarly engaging activity, then sup-
port is lent to theories of more “direct” effects of the
music on cognitive processes (see Leng & Shaw, 1991;
and “Neuronal Network Priming”).
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Neurobiological Mechanisms
Underlying the Effect of Music on
Cognitive Performance in Humans
The claim that music enhances cognitive perfor-
mance can also be explored via an examination of poten-
tial mediators. If music can be shown to influence physi-
ological or psychological mechanisms that are known to
affect cognitive performance, then it can be inferred
that music has at least the potential to influence cogni-
tive performance. Although there are numerous psycho-
logical explanations for this relationship, for instance,
exposure to music might focus attention, reduce distrac-
tion, or enhance motivation for performing a certain
task, a primarily neurobiological approach will be taken
to address this question here.
Affective Mediation
One of the most defining features of music is its capac-
ity to induce strong emotions in listeners (Abeles &
Chung, 1996; Gabrielsson, 2001), the impact of which is
perceived both physically and mentally (Panzarella,
1980). The central method of expression in music tends
to be affective rather than referential, and modulation
or experience of emotion is one of the primary reasons
people give for listening to music (Konecni, 1982;
Sloboda, 1992; Sloboda & O’Neill, 2001). Whereas
Tolstoy labeled music “the shorthand of emotion,”
cognitivists deny that music actually induces emotions in
listeners (Kivy, 1989, 1990; Langer, 1942). The evidence
from lesion, imaging, EEG, and dichotic listening tasks,
however, challenges this position, suggesting that musi-
cally induced emotions are real and processed in a simi-
lar way to nonmusical emotions. In particular, the right
hemisphere appears to play a greater role than the left
hemisphere in the interpretation of emotions in music
(Blood, Zatorre, Bermudez, & Evans, 1999) as observed
with nonmusic emotions (e.g., Adolphs, Damasio,
Tranel, & Damasio, 1996; Blonder, Bowers, & Heilman,
1991; Ley & Bryden, 1979, 1982; McKeever & Dixon,
1981; Safer & Leventhal, 1977; Tucker, Watson, &
Heilman, 1977). As with processing of nonmusical emo-
tions (Gainotti, 1972; Silberman & Weingarter, 1986),
there are also data indicating that lateralization of pro-
cessing musical emotions may depend on valence
(Altenmüller, Schürmann, Lim, & Parlitz, 2002; Blood &
Zatorre, 2001; Gagnon & Peretz, 2000; Schmidt &
Trainor, 2001; Tsang, Trainor, Santesso, Tasker, &
Schmidt, 2001), particularly when participants are asked
to focus on the experienced feeling evoked by the music
(Blood & Zatorre, 2001; Schmidt & Trainor, 2001).
If music enhances cognitive performance via affective
modification, then concomitant changes should also be
 Rickard et al. / FACILITATORY EFFECTS OF MUSIC ON COGNITIVE PERFORMANCE
observed in autonomic indices of arousal (Rickard,
2004; Scherer & Zentner, 2001; Trainor & Schmidt,
2003). Arousal is a primary dimension of emotion in
both nonmusical (Russell, 1979, 1980) and musical emo-
tions (North & Hargreaves, 1997; Ritossa & Rickard,
2004; Schubert, 1999). The effect of emotional arousal
on cognitive performance has been well established in
the nonmusical domain (e.g., Cahill, Gorski, & Le, 2003;
Cahill & McGaugh, 1998; Cahill, Prins, Weber, &
McGaugh, 1994; Hamann, Ely, Grafton, & Kilts, 1999;
O’Carroll, Drysdale, Cahill, Shajahan, & Ebmeier, 1999;
Quevedo et al., 2003). For example, recall of material is
significantly improved when associated with emotional
arousal in both normal (Cahill et al., 2003) and memory-
impaired participants (Kazui et al., 2000). The level of
arousal at both the time of learning (e.g., Humphreys &
Revelle, 1984) and retrieval (Eysenck, 1976) is thought
to be critical in memory processing. Schellenberg and
colleagues convincingly demonstrated that improve-
ments in spatial temporal performance following passive
exposure to music (see “Spatial Performance” section)
are likely to be attributable to listeners’ enjoyment of the
piece, and thus mediated by increased arousal (Husain
et al., 2002; Nantais & Schellenberg, 1999; Schellenberg,
2001; Schellenberg & Hallam, in press; Schellenberg,
Nakata, et al., in press; Thompson et al., 2001). It is possi-
ble then that the increased physiological arousal associ-
ated with experiencing emotions is responsible for any
effects of music on cognitive performance.
Autonomic nervous system responses. Numerous studies
have demonstrated autonomic nervous system re-
sponses to a range of emotion-inducing pieces of music
(see Bartlett, 1996, for review). Skin conductance, in
particular, appears to be a useful indicator of musically
induced emotional arousal (Khalfa, Peretz, Blondin, &
Manon, 2002; Rickard, 2004; Vanderark & Ely, 1992,
1993). Despite some promising findings, however, the
weight of contradictory or null findings is of concern.
Given the limited sensitivity of autonomic indices as
measures of emotion, particularly because many studies
average responses over time, it is not surprising that the
impact of extraneous variables might often be larger
than that of the music. On the other hand, owing to their
immediacy, these indices are quite specific to emotional
responses. This is in contrast to measurement of emo-
tional processing in the brain, which is subject to greater
interference from nonaffective processing, such as theo-
retical analysis of the music or recall of associations with
the piece (Peretz, 2001).
There is still a clear need for improved methodology
in this area of research. One solution has been to mea-
sure extraneous variables (personal, musical, and situa-
tional) that may interact with music to influence physio-
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241
logical responses (Abeles & Chung, 1996; Gabrielsson,
2001; McMullen, 1996). This has been attempted in iso-
lated cases, for instance, with regard to personality
(Crawford & Strapp, 1994; Furnham & Allass, 1999; Ries,
1969), musical training (Vanderark & Ely, 1992), and
features of the music itself (Panksepp, 1995; Sloboda,
1991). Sloboda, for instance, lists melodic appoggiaturas
(or “leaning notes”) and new harmonies as important
predictors of emotional response in listeners. The prac-
ticality, however, of measuring an adequate range of
such variables is countered by the sample size that would
be required for meaningful statistical analysis. A second
approach to limiting the impact of the overwhelming
variability between listeners is to enhance the signal-to-
noise ratio, so that the emotional response to music is
salient among other responses. Utilizing pieces of music
selected by the participants rather than experimenters
increases the likelihood that the participant will elicit
an intense emotional response that may be sufficiently
marked to stand out, despite interactions with other vari-
ables. This approach has been used with some success in
a limited number of studies. Rickard (2004), for
instance, found significant increases in skin conduc-
tance and number of chills in response to participant-
selected “emotionally powerful” pieces. Much smaller
increases in these measures were observed in response
to a nonemotional, but arousing, music piece or an emo-
tionally powerful nonmusical piece (film scene), sug-
gesting that music may indeed be an extremely powerful
elicitor of emotions. “Chills” (or goosebumps, an index
of autonomic arousal) appear to be robustly elicited
when the methodology enables participants to select
their own piece of music (Blood & Zatorre, 2001;
Gabrielsson, 2001; Goldstein, 1980; Panksepp, 1995;
Rickard, 2004). On the other hand, the use of participant-
selected music greatly reduces the opportunity to
observe the impact of various music elements in any sys-
tematic way, because the combination of these elements
will differ widely across participants.
A third approach is to explore a wide range of auto-
nomic indices across a great number of musical stimuli
varying in affective tone. Using this methodology,
Krumhansl (1997) observed a different pattern of auto-
nomic responses depending on the type of emotional
response elicited by the music. She found that “sad”
musical excerpts were associated with decreases in
skin conductance level, increased blood pressure,
and slower heart rate, whereas “happy” pieces caused
greater changes in respiration measures. Importantly,
Krumhansl monitored physiological responses to music
excerpts continuously (second-by-second), which is an
additional means of enhancing data accuracy (see also
Harrer & Harrer, 1977). Nyklicek, Thayer, and Van
242 BEHAVIORAL AND COGNITIVE NEUROSCIENCE REVIEWS
Doornen (1997) also found that “happy,” “sad,”
“serene,” and “agitated” music could be differentiated
by respiratory and chronotropic indices. These “auto-
nomic signatures” (Trainor & Schmidt, 2003) are consis-
tent with findings from research with nonmusical emo-
tional stimuli, where sympathetic nervous system indices
such as heart rate and finger temperature have been
found to differentiate the primary emotions in facial ex-
pressions to some extent (Levenson, Ekman, & Friesen,
1990). Khalfa et al. (2002) demonstrated greater
increases in skin conductance in participants exposed to
“fearful” or “happy” musical excerpts than to “sad” or
“peaceful” excerpts, which appears to confirm the sug-
gestion from nonmusic emotion literature that skin con-
ductance is an index of arousal, rather than valence
(Bradley & Lang, 2000). There is also some consistency
with the actual pattern of changes across musical and
nonemotional stimuli, with differences potentially
attributable to methodological issues (see Krumhansl,
1997).
Neurochemical responses. Emotional arousal is accompa-
nied by the release of a number of neurochemicals,
which are known to moderate cognitive performance.
These include transmitters and hormones involved in
generalized arousal control systems, such as the cate-
cholamines noradrenaline and adrenaline, the hor-
mones cortisol and ACTH, and neurotransmitters
involved in reward and pleasure (e.g., Ashby, Isen, &
Turken, 1999; Cahill et al., 1994; Gibbs & Summers,
2002; McIntyre, Hatfield, & McGaugh, 2002; O’Carroll
et al., 1999; Sara, 1991). Music may therefore affect cog-
nitive performance via neurochemical changes that
result from an emotional response. For example, dopa-
mine and beta-endorphin have been implicated in
responses to music (Hodges, 1996; Panksepp &
Bernatzky, 2002). However, empirically derived evi-
dence that music can stimulate release or inhibition of
these chemicals is currently limited, partly because of
ethical constraints preventing psychopharmacological
manipulations with humans. Preliminary data, however,
suggest that administration of the opiate receptor antag-
onist naloxone reduces the number of chills experi-
enced by listeners in response to emotionally powerful
music (Goldstein, 1980). This suggests that endorphins
may play an important role in the strong emotional
response to music. Interestingly, Harrer and Harrer
(1977) found that participants retained their emotional
responses to music following administration of a moder-
ate dose of tranquilizers that completely suppressed
autonomic responses to music. This is inconsistent with
the considerable literature suggesting that autonomic
arousal underlies intensity of the emotional response to
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music. However, because the detail of this component of
Harrer and Harrer’s research is minimal, it is difficult to
evaluate its conclusiveness.
A number of studies have measured transmitters or
their metabolites in urine, blood, or saliva during or fol-
lowing music exposure (e.g., Bartlett, Kaufman, &
Smeltekop, 1993; Möckel et al., 1994; see Bartlett, 1996,
for a review). However, few have isolated the neuro-
chemical responses underpinning the emotional re-
sponse to music. Rickard (2004) measured salivary
cortisol in response to emotionally powerful music but
found no significant difference when compared with
relaxing or arousing music. Vanderark and Ely (1992,
1993) measured noradrenaline, adrenaline, and cortisol
responses following exposure to “joyful” (Jupiter from
Holst’s The Planets) or “horrific” (Penderecki’s Threnody
for the Victims of Hiroshima) music. Although the format in
which these studies is reported is somewhat incomplete,
the overall results appear to suggest that music produced
significant decreases in plasma noradrenaline, regard-
less of the type of music presented or participants’ musi-
cal background. In contrast, both types of music pro-
duced increases in cortisol in musically trained
participants only. Hirokawa and Ohira (2003) compared
noradrenaline responses to music that was described by
participants as having high or low “uplifting” qualities
and found no difference. Gerra et al. (1998) found that
although exposure to classical music enhanced partici-
pants’ emotional state, there was no associated change in
beta-endorphin, noradrenaline, or cortisol levels. In
contrast, exposure to “techno” music was associated with
worsening of emotional state and an increase in beta
endorphin, noradrenaline, and cortisol. In summary,
the available data do not provide sufficient evidence of a
consistent change in the release of any neurochemicals
following exposure to emotion-inducing music. Never-
theless, the methodology is promising and the results to
date suggest that noradrenaline, dopamine, and cortisol
are all worthy candidates for future research.
Nonaffective Mediation
Neuronal network priming. Extensive research on the
perception of music suggests that there are specialized
brain structures (modularity) for music processing (see
Altenmüller, 2003; Peretz & Coltheart, 2003). A second
major hypothesis as to how music might enhance cogni-
tive performance is by direct activation of similar
neuronal networks (independent of emotional
response). This mechanism is consistent with the con-
cept of “priming,” in which exposure to one type of
material improves learning or identification of another
related type of material (Tulving & Schachter, 1990). For
instance, inducing positive affect can enhance recall of
 Rickard et al. / FACILITATORY EFFECTS OF MUSIC ON COGNITIVE PERFORMANCE
“happy” information (Bower, 1981; Nasby & Yando,
1982; Teasdale & Fogarty, 1979). However, the stimuli to
be recalled do not have to be affective in nature, and
there is considerable evidence for priming of nonaffec-
tive visual or verbal material (see Schellenberg, 2001).
This framework has been applied to music-related
effects in an attempt to explain how exposure to music
might enhance performance on a cognitive task.
Rauscher et al. (1995) argued that the short-term
increase in spatial-temporal scores observed following
exposure to 10 min of a Mozart sonata was due to coinci-
dent activation of neuronal “trions”; “musical activity
strengthens inherent neural firing patterns that are also
utilized by spatial-temporal tasks” (p. 427). In other
words, prior exposure to complex music “primed” the
neuronal networks that would also be recruited to per-
form the spatial-temporal mental manipulations
required to complete the Paper Folding and Cutting
(PFC) task from the Stanford-Binet IQ test.
Despite a rather abstruse presentation, there is some
gross appeal in this hypothesis. It has been established
that the brain can be permanently modified by music
training (e.g., see “Synaptic Plasticity” section), so short-
term plasticity in response to passive exposure is not
unreasonable. The concept is also reminiscent of Hebb’s
“reverberating” circuits, in which neuronal connections
are temporarily strengthened by learning. Schellenberg
(2001), however, discounts priming (or the related pro-
cess “transfer” of learning) as a likely explanation for the
Mozart effect. He argues that music and spatial-temporal
performance are too dissimilar. In particular, although
cross-modal priming is known to occur, instances
whereby pre-exposure to auditory stimuli primes recog-
nition of visual stimuli have not been demonstrated (see
Husain et al., 2002). Nonetheless, enhancement of ver-
bal learning has previously been observed following
music (Chan, Ho, & Cheung, 1998; Morton et al., 1990),
despite the dissimilarity of these two stimuli types. Fur-
thermore, there is a range of skills that are arguably
involved in processing of both music and spatial-
temporal tasks, including abstraction and transforma-
tion (Serafine, 1988). Rauscher et al. (1995) argued that
when listening to Mozart’s sonata, the brain must pro-
cess the pattern of spatial intervals as they change over
time; this type of mental manipulation bears some simi-
larity to that performed in the PFC task. In support,
Rideout and Laubach (1996) found that music exposure
enhanced spatial temporal performance and that im-
proved performance correlated significantly with
increases in EEG power recorded in temporal and fron-
tal brain regions during task performance. Sarnthein
et al. (1997) explored EEG coherence patterns during
performance of a spatial-temporal task and during expo-
sure to the Mozart sonata. They then measured coher-
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243
ence patterns during spatial performance preceded by
exposure to Mozart and examined the difference in pat-
terns during the two task performances. In 3 of 7 partici-
pants, a pattern of coherence involving prefrontal and
temporo-parietal regions was present both during music
exposure and during task performance that followed
music exposure. They concluded that the coherent activ-
ity induced by music extended several minutes and was
even “carried over” into the performance of the spatial
temporal task. Theoretically, outright rejection of the
neuronal network priming hypothesis may therefore be
premature. Although at a cognitive level, preferred
music (or other enjoyed stimuli) and a spatial task do not
appear to be associated, at a neurophysiological level,
they may well be.
Synaptic plasticity. The mechanisms underlying any
longer term effect of music exposure (particularly music
training) are of considerable interest. However, to date,
such mechanisms have not been widely sought within
music neuroscience research. Many skills and experi-
ences are associated with music training, including time
spent with a nonparental adult, disciplined practice, fine
tuning of sensorimotor coordination, extensive
memorization and sustained attention, spatial-temporal
visualization, pride in achievements and performance,
exploration of emotions, and team work (Schellenberg,
2001). Any of these could transfer to benefits in other
aspects of cognitive performance. Music training also
develops pitch and rhythm discrimination, each of
which has been claimed to benefit reading performance
(Douglas & Willats, 1994; Lamb & Gregory, 1993).
Music may also provide environmental enrichment.
Nonmusic research has established that environmental
enrichment has significant and long-lasting effects on
the brain, increasing the potential for future learning via
increased neuronal growth (Rosenzweig, 1984) or
enhanced cortical maturation (Raine et al., 2001). Con-
sistent with this possibility, music training has been
found to modify brain plasticity. For instance, areas of
the cortex associated with finger control are larger in
string players (Elbert, Pantev, Wienbruch, Rockstroh, &
Taub, 1995) and the size of the corpus callosum is larger
in musicians, particularly those who began lessons be-
fore 7 years (Schlaug, Jancke, Huang, & Steinmetz,
1995). It is possible that either of these concomitants
(associated skill enhancement or environmental enrich-
ment) could be responsible for improvement in other
cognitive functions. Nevertheless, further systematic and
well-controlled research is required to establish whether
music training does in fact have any positive effect on
academic or other cognitive performance. Unfortu-
nately, it is difficult to examine changes in synaptic plas-
ticity at the neuronal level in humans, although postmor-
244 BEHAVIORAL AND COGNITIVE NEUROSCIENCE REVIEWS
tem examination of acoustically enriched compared to
“normal” brains would be informative in this respect.
Summary and Additional Limitations
of Human Studies
In the section “Effect of Music on Cognitive Perfor-
mance in Humans,” evidence of music’s impact on cog-
nitive performance was reviewed. Although there is
some evidence that music might improve spatial perfor-
mance and learning, it must be concluded that the lit-
erature on the effects of music on physiology and cogni-
tion remains characterized by its inconsistencies (see
Bartlett, 1996; Schellenberg, 2001). There is little defini-
tive evidence that music enhances cognitive perfor-
mance in humans. The lack of conviction in the data
might simply indicate that cognitive performance can-
not be improved by musical treatments. However, in the
section “Neurobiological Mechanisms Underlying the
Effect of Music on Cognitive Performance in Humans,”
several means by which music could have a cognitive-
enhancing effect were reviewed. A strong candidate is
emotion induction, which subsequently increases physi-
ological arousal. Evidence is accumulating that release
of arousal-related neurochemicals (such as catechola-
mines and corticosteroids)—which are believed to
mediate enhancements in cognitive performance—also
accompanies listening to music.
Given the potential of music to stimulate emotions,
physiological arousal, and shared neuronal networks, it
would be surprising if music did not influence cognitive
performance, if only transiently. It is possible then that
the effects of music on cognitive performance are often
masked by methodological problems. It has been argued
that inconsistencies in research findings are partly due
to the difficulty in controlling, or accounting for, all
extraneous variables that could potentially moderate or
confound the results (Hallam et al., 2002; Spintge &
Droh, 1992). However, there are other realities of hu-
man research that also limit the extent to which con-
clusions can be drawn from these data.
First, it is all but impossible to obtain participants who
have had no prior exposure to music. Human fetuses
have been shown to respond to music from as early as the
second trimester (Gagnon, Hunse, Carmichael, Fellows,
& Patrick, 1987), and babies are regularly exposed to
hummed tunes and lullabies in all cultures. It will there-
fore always be difficult to attribute any effects of music to
the music itself, rather than to related factors such as
stimulus familiarity or preference. Second, many tech-
niques exploring underlying mechanisms are corre-
lational or indirect. For instance, although studies dem-
onstrating that similar brain regions are activated by
music and cognitive performance are intriguing, causal
inferences can only be made with caution. It remains dif-
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ficult to dissociate specific from nonspecific (such as
attention and retrieval) effects, or to conclusively dem-
onstrate the direction of causality. The lack of temporal
resolution in popular imaging techniques (such as PET)
may also dilute significant effects. On the other hand,
EEG has poor spatial resolution and is of limited use for
measuring subcortical activation (unless depth elec-
trodes are used). More direct neurophysiological or
neurobiological examination of responses to music is
used less often in human participants. When such mea-
sures are obtained (for instance, salivary cortisol levels
are measured or opioid antagonists administered), the
data are fascinating but still limited by confounds and
variability, which are difficult to exclude. In the next sec-
tion, we explore whether some of the methodological
constraints of human music psychology research can be
overcome by using animal models.
ANIMAL STUDIES
Are Animal Models Appropriate for
Music Psychology Research?
The call to explore the effects of music on physiology,
emotion, and cognition via animal studies is not entirely
novel. A number of prominent researchers have argued
that there are compelling reasons for this approach
because there are many similarities in how animals and
humans process complex sounds (Hodges, 1996;
Mikiten, 1996; Panksepp & Bernatzky, 2002). In the fol-
lowing sections, we review what has been learned about
the effects of music on cognitive performance via non-
human animal (hereafter referred to as “animal”)
research. Such findings should not be interpreted with-
out first having considered the utility of animals as mod-
els of human music experience.
The Utility of Animal Models
In comparison to human research, adherence to
systematic experimental design is generally easier in
animal research. Animals can be randomly allocated to
appropriate experimental and control groups with less
concern for time or repetition demanded of participants
(although this should still be constrained by ethical prin-
ciples of refinement, replacement, and reduction of ani-
mal use) and superior control of extraneous variables,
such as testing environment and time of testing. Rightly
or not, more direct methods (psychopharmacological
experiments, invasive recording techniques, and experi-
mental lesions) are often used with animal subjects that
are typically prohibited or impractical with human par-
ticipants. If required, animals can be isolated entirely
from music exposure (including prenatally), or the
music to which they are exposed can be systematically
controlled, enabling some distinction of innate, as
 Rickard et al. / FACILITATORY EFFECTS OF MUSIC ON COGNITIVE PERFORMANCE
opposed to learned, “musical” faculties (Hauser &
McDermott, 2003). Within-group variation is also con-
siderably reduced, in that individual musical prefer-
ences, tastes, musical training/knowledge, and associa-
tions with particular pieces of music, personality, and so
on are all but excluded in animal species. The auditory
stimulus can be more easily deconstructed to ascertain
the effects of stimuli that vary only on pitch, amplitude,
timbre, rhythm, meter, harmony, scale, or form (Hulse &
Page, 1988). Although such oddities may introduce a
new batch of confounding variables in human partici-
pants (such as novelty, dislike, and boredom), these fac-
tors are less likely to influence animal responses to audi-
tory stimuli because animals have not typically become
acculturated to more complex music. This methodology
enables considerably more rigorous, systematic, and
insightful studies to be performed than is generally
feasible with humans.
Many will argue that by excluding these characteris-
tics, the essence of music is lost; that is, the richness of the
music experience depends on these socioemotional fac-
tors. However, the remnants once these factors have
been excluded can be equally informative—these funda-
mental responses that we share with animals arguably
form the foundation of our elementary responses to
music (see “Input From Comparative Psychology”
section). The auditory elements that produce such
responses in animals may be the ultimate music “univer-
sals” (see Hulse & Page, 1988), common across all
human cultures, all subpopulations and ages, and pres-
ent even in compromised individuals (due to illness,
dementia, and perhaps even coma; e.g., Hughes,
Daaboul, Fino, & Shaw, 1998; Swartz, Hantz, Crummer,
Walton, & Frisina, 1989).
Input From Comparative Psychology
An assumption implicit in animal research in this con-
text is that there is some level on which animal responses
can be generalized back to human responses to music.
For this to be possible, there must be some commonality
in how animals and humans react to musical stimuli.
Although any similarity of animal calls and songs to
human music is likely to be coincidental (Hauser &
Konishi, 1999; Hauser & McDermott, 2003; Wallin et al.,
2000), there is good reason to believe homologies exist
across humans and other animals in the perception of
musical sounds. In the remainder of this section, we
review the research on animal perception of human
music. The focus will be on identifying which musical
faculties animals do have, such that there can be clear
demarcation with regard to the scope of comparison
from animals to humans. Musical faculties found in ani-
mals indicate that the underlying brain substrates are
unlikely to have adapted specifically for music. Instead,
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245
these brain areas probably subserve more generalized
auditory functions (Hauser & McDermott, 2003;
Panksepp & Bernatzky, 2002; Weinberger, 1999). For
instance, it is well established that prosody in both ani-
mal and human vocalizations reflects the communica-
tor’s emotional state. Panksepp and Bernatzky (2002)
suggested that music may have developed from the
“intrinsic emotional sounds we make” (p. 134). Animals
also appear to use tempo, pitch variations, accents, and
so forth to communicate fear, anger, and happiness. It
has been argued that humans probably utilized this
capacity of the brain for music in a similar way as
nonlanguage areas of the brain are thought to have been
utilized for language (Hauser & McDermott, 2003;
Weinberger, 1999). Despite animals being nonlinguistic,
several mechanisms underlying speech perception are
shared by animals (Hauser, Chomsky, & Fitch, 2002;
Kuhl, 1986), allowing much to be learned about human
language from animals. The elements that are not found
in nonhuman species, of course, also tell us much about
“what makes human music perception unique” (Hulse &
Page, 1988, p. 427).
Sensation and perception. First we will focus on sensitiv-
ity and then on perception and discrimination. Sensitiv-
ity: Hearing range (the sensation of sound) varies enor-
mously across species and can be defined in terms of
frequency, sound pressure, and duration. Although
hearing range does not dictate whether an animal brain
has music-processing capabilities, it does constrain ex-
perimental design and interpretations that can be drawn
from animal research, given that human music is
designed for human ears. Humans are able to hear fre-
quencies ranging from approximately 20 Hz (which is
very close to the frequency of the lowest note on a piano
keyboard) up to 20 kHz. However, humans are most sen-
sitive to frequencies between 200 Hz and 8 kHz, particu-
larly those between 2 and 4 kHz (Jackson, Heffner, &
Heffner, 1999). The frequency range of our greatest sen-
sitivity (around 4 kHz) corresponds to the top octave of
the piano keyboard, and frequencies above this range
are not typically perceived as “musical” pitches
(Carterette & Kendall, 1999). The lowest perceptible
sound is typically 0 decibels, and 140 dB sounds are pain-
ful (e.g., jet airplane taking off). The shortest duration of
sounds we can usually detect is approximately 100 ms
(Krumhansl, 2000).
Although there is considerable overlap between the
hearing ranges of many animals and humans, animal
audiograms often reveal differences in peak sensitivity
and frequency thresholds. Rodents’ audiograms, for
example, are shifted considerably to the right, with labo-
ratory rats’ greatest sensitivity between 8 and 32 kHz
(Heffner, Heffner, Contos, & Ott, 1994). Rhesus mon-
keys have similar sensitivity to humans, whereas chim-
246 BEHAVIORAL AND COGNITIVE NEUROSCIENCE REVIEWS
panzees have a narrower range of sensitivity. Dogs tend
to have much broader ranges of sensitivity (Sales & Pye,
1974; Warfield, 1973). The hearing range of many birds
is quite comparable to humans, ranging from 100 Hz to
29 kHz (depending on the species), with peak sensitivi-
ties between 2 and 4 kHz (although sound pressure lev-
els may need to be somewhat higher than for humans for
these frequencies to be heard). Many animals are, how-
ever, more sensitive to lower sound pressures than are
humans and may have greater temporal resolution. For
instance, birds can discriminate notes lasting only 5 ms
(20 times shorter than humans), and only a few millisec-
onds apart, whereas humans discriminate sounds when
between only 12 and 50 ms apart (Wilkinson & Howse,
1975).
Perception and discrimination: There is no doubt that
animals are able to perceive complex species-specific
vocalizations (e.g., Dooling, 1989). However, it is diffi-
cult to assess whether animals perceive musical sounds in
a similar way as do humans. Weinberger and McKenna
(1988) demonstrated that the auditory neurons of anes-
thetized cats responded to tonal contour. Even so, this
direct neurophysiological approach has not been
explored sufficiently to provide further insight into this
issue. One method that has been illuminating in this
respect is the examination of perceptual invariance (see
Carterette & Kendall, 1999; Hulse & Page, 1988; Hulse,
Takeuchi, & Braaten, 1992, for reviews). Perceptual
invariance refers to a structural characteristic that is per-
ceived as the same even when transformed. For instance,
a melody is perceived by humans to be the same even
when transposed into another key or up several octaves
(Deutsch, 1999; Dowling & Harwood, 1986). This is
because we encode the pitch intervallic relations (or
melodic contour) rather than the absolute pitches them-
selves. Humans demonstrate perceptual invariance for
frequency structures (pitch, melody, harmony, and tim-
bre), intensity (loudness), and time structures (rhythm
and meter) (see Hulse et al., 1992, for review). If animals
also demonstrate perceptual invariance for these musi-
cal structures, then there is a prima facie case that animals
have the necessary perceptual apparatus for hearing
music.
The most detailed investigations of perceptual
invariance have occurred with the European starling
songbird. A classic operant task is used in which the birds
peck at different disks to demonstrate discrimination of
presented sounds. Using this methodology, Hulse and
colleagues have demonstrated that starlings are able to
discriminate different timbres (Braaten & Hulse, 1991),
intensities (Bernard & Hulse, 1992), rhythms (Hulse,
Humpal, & Cynx, 1984), pitches, and pitch patterns
(Hulse, 1989) and perceive the missing fundamental in
harmonic complexes (Cynx & Shapiro, 1986). However,
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Hulse and colleagues found that, with regard to pitch
processing, starlings could not discriminate when the
pitches were transposed beyond the range used in train-
ing, nor could they generalize recognition to notes an
octave apart. This suggests that birds rely on local cues
rather than melodic contour to perform pitch-related
discrimination. It has been concluded that, unlike
humans, birds tend to rely on absolute pitch processing
(or perfect pitch) more than relative pitch processing
(i.e., the pattern of intervals between pitches).
Nonhuman primates, our closest ancestors, also fail
to demonstrate perceptual invariance for melodies.
Monkeys can discriminate complex acoustic patterns
but do so by focusing on local pitch features (e.g., the last
two notes), rather than the entire melody (D’Amato,
1988; D’Amato & Salmon, 1984). This discrimination is
on the basis of absolute rather than relative pitch (Hulse
et al., 1992). When transposed to new frequencies, or
even an entire octave, monkeys no longer recognize the
melody (D’Amato, 1988). In sum, animals appear to per-
ceive perceptual invariance in the majority of musical
structures, except melody.
Despite a preference for utilizing absolute pitch, star-
lings appear to be capable of processing relative pitch
information as well as absolute pitch in their initial learn-
ing (MacDougall-Shackleton & Hulse, 1996). Demon-
stration of relative pitch knowledge in starlings requires
specific training, which is not true in human adults. Simi-
larly, monkeys can be trained to utilize relative pitch if
absolute pitch cues are not available (Brosch, Selezneva,
Bucks, & Scheich, 2004). It is therefore also of interest
that there is some evidence that children (3 to 5 years of
age) are better at absolute than relative pitch processing
(e.g., Saffran & Griepentrog, 2001; White, Dale, &
Carlsen, 1990; see also Hulse et al., 1992). However, con-
flicting evidence (Trehub, Bull, & Thorpe, 1984;
Trehub, Thorpe, & Morrongiello, 1987) indicates that
infants preferentially use relative pitch processing.
Regardless, children’s capacity for using relative pitch
appears to improve with training (e.g., Sergeant &
Roche, 1973), which is not entirely inconsistent with the
findings in starlings.
Indeed, several studies challenge the claim that ani-
mals do not demonstrate perceptual invariance for mel-
odies. Ratcliffe and Weisman (1985) reported that chick-
adees occasionally shift their two-tone song in a ratio
manner, suggesting relative pitch. Gerbils can discrimi-
nate rising from falling frequency-modulated tones
(Wetzel, Wagner, Ohl, & Scheich, 1998). Interestingly,
right hemisphere lesions impair this discrimination,
whereas no impairment is observed with left hemisphere
lesions (Wetzel, Ohl, Wagner, & Scheich, 1998). The
underlying neural substrate required for this task in the
gerbil is not known; however, it is intriguing that right
 Rickard et al. / FACILITATORY EFFECTS OF MUSIC ON COGNITIVE PERFORMANCE
hemisphere lesions also impair melody discrimination
(Zatorre, 1985) and interpretation of emotions in music
in humans (Blood et al., 1999; Blood & Zatorre, 2001;
Bryden, Ley, & Sugarman, 1982). Moreover, there are
several findings that suggest that at least some mammals
may demonstrate octave equivalence. Bottlenose dol-
phins have been heard to spontaneously shift pitches an
octave in their production of a learned pitch pattern
(Richards, Wolz, & Herman, 1984). Rats have also been
claimed to show octave generalization of a single pitch
(Blackwell & Schlosberg, 1943), although the method-
ology of this very early study may not have been that
rigorous.
Most intriguingly, Wright, Rivera, Hulse, Shyan, and
Neiworth (2000) found evidence of octave generaliza-
tion of melodies in rhesus monkeys. These monkeys
judged a melody as the same when transposed by one or
two octaves. This indicates the use of relative pitch,
although to a lesser extent than humans because the
monkeys did not recognize a melody as the same when it
was shifted to a different key (that is, less than an octave
translation). Furthermore, this effect was only observed
when tonal melodies were used (i.e., consisting of tones
from the diatonic scale), but not when random-synthetic
melodies or atonal melodies were used. This raises the
possibility that the monkey’s brain is innately wired to
respond to tonal melodies. Rats (Cross, Halcomb, &
Matter, 1967) and birds (Watanabe & Nemoto, 1998)
have also been reported to show a preference for clas-
sical over atonal music (see “Affective Responses to
Music?” section). This claim is quite extraordinary and
would strongly justify generalization of findings on ani-
mal responses to music to humans. However, in addition
to the use of small sample sizes (e.g., two in the study by
Wright et al., 2000), it is possible that these animals had
been exposed to Western music and thus learned our
tonal system. Indeed, McDermott and Hauser (2004)
ensured that five cottontop tamarins were not exposed
to any music prior to testing and found that the monkeys
showed no preference for consonant over dissonant
melodies, casting doubt on animal perception of the
human tonal system. It is interesting to note, however,
that both birds (Hulse, Bernard, & Braaten, 1995) and
monkeys (Izumi, 2000) can be trained to discriminate
consonant from dissonant intervals.
Birds also appear capable of discriminating between
larger scale musical characteristics. Porter and Neuringer
(1984) trained pigeons to discriminate between 1 min
excerpts from a piece by Bach from 1 min excerpts by
Hindemith, using a peck-disk reinforcement schedule
procedure. This discriminative ability could not be
attributed to any single note or phrase in the pieces,
because it persisted even when the 1 min excerpts were
drawn randomly from 20 min excerpts. In a further
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247
study, they found that 4 out of the 5 pigeons tested could
categorize novel excerpts by Baroque and 20th century
composers as belonging to either the “Bach” category or
the “Stravinsky” category. This apparent ability to per-
ceive musical genre was also observed in Java sparrows
(Watanabe & Sato, 1999). Can we conclude from such
findings that animals can also perceive musical styles or
form? It is difficult to fully interpret these findings be-
cause there are many music features (such as timbre and
tempo) that would differ across genres, any one of which
the animals may have been utilizing for discrimination.
Nevertheless, the capacity of some bird species indi-
cates that there may be greater similarity in the way that
animals and humans hear human music than is often
assumed.
Affective responses to music? In the absence of subjective
report, behavioral and physiological concomitants of
human affective responses have been used to infer affec-
tive state in animals. Much of the relevant research has
been performed in the context of environmental enrich-
ment for agricultural or laboratory animals, although
anecdotal claims of companion animals “enjoying”
human music are abundant. Although such investiga-
tion has been criticized as anthropomorphic in that it
assumes animals “hear” music in the same way as humans
do (see Newberry, 1995), the above review of animal
capabilities suggests that many animals share at least
basic perceptual abilities with humans. Given their past
experience and associations with music, personalities,
and cultural embeddedness, humans will have addi-
tional affective responses to music over and above those
of animals. Nevertheless, given that (a) at least some
aspects of emotion are governed by evolutionarily primi-
tive parts of the brain that are functionally homolo-
gous to those in other nonhuman mammalian brains
(Panksepp, 2003; Panksepp & Bernatzky, 2002) and (b)
affective responses may prove to be the major mediator
of music’s facilitatory effect on cognitive performance in
humans (see “Affective Mediation” section), it is clearly
of interest to explore whether animals demonstrate any
affective response to music. In the context of emotional
syntax theories of music, it may even be useful to con-
sider animals’ affective responses to music as the most
basic of the bottom-up processes, which in humans are
described as “automatic, unconscious, preprogrammed,
[and] ‘brute’” (Narmour, 1991, p. 3). A propensity for
such responses does not seem unreasonable given ani-
mals’ perceptual capacity is subject to some of the same
basic gestalt principles that can manipulate expectation
and confirmation (or violation of), and thus emotional
response, in humans.
A commonly used measure of affective response in
human listeners is preference (Berlyne, 1971; Konecni,
1979; Miller, 1992; North & Hargreaves, 1997). Prefer-
248 BEHAVIORAL AND COGNITIVE NEUROSCIENCE REVIEWS
ence for auditory stimuli can be directly measured in ani-
mals using an apparatus in which the animal can control
the onset of different sounds (Blom et al., 1992; Park &
Balaban, 1991). In a preliminary study, two javabirds
that were given the choice of three perches spent signifi-
cantly more time on a perch that triggered music by
Bach than on perches that triggered music by Schoen-
berg or silence (Watanabe & Nemoto, 1998). (The
music piece triggered by each perch was reversed
between trials to control for nonspecific factors that
could influence preference.) These same birds also
spent more time on a perch that triggered music by
Bach’s contemporary Vivaldi than on a perch that trig-
gered music by Schoenberg’s contemporary Carter, sug-
gesting that the birds preferred classical to 20th century
music. Rats also appear to imprint more easily to classical
than atonal music. When given a choice between com-
partments that triggered music by either Mozart or
Schoenberg, rats that had been reared on Mozart’s
music spent more time in the former compartment. In
contrast, rats reared on Schoenberg’s music showed no
compartment preference (Cross et al., 1967). The
authors suggested that Schoenberg’s atonal music may
be too complex for rats to form an attachment. Fi-
nally, although nonhuman primates demonstrated no
preference for consonant over dissonant intervals
(McDermott & Hauser, 2004), preliminary findings indi-
cate that they may show a preference for slow over fast
stimuli (60 vs. 400 clicks/min) and for lullabies over fast
technomusic (McDermott, 2005). It is of note, however,
that the sample sizes in these studies were exceptionally
small (4 in Watanabe & Nemoto, 1998, and 5 in
McDermott & Hauser, 2004) and that only 2 of the java-
birds in the former study showed a preference for the
music (the other 2 showed no preference for any perch).
Thus, replication is required to ensure these results did
not occur by chance.
Preference can also be implied via certain behaviors,
such as increased approach or social activity. Exposure to
classical music has been claimed to increase the growth
rate and/or feeding behavior in a range of species,
including poultry (Gvaryahu, Cunningham, & van
Tienhoven, 1989; Ladd, Albright, Beck, & Ladd, 1992),
cattle (Zhuchzhu & Andonov, 1995), horses (Houpt,
Marrow, & Seeliger, 2000), and fish (Vasantha,
Jeyakumar, & Pitchai, 2003), although further replica-
tion is still required. Country music exposure enhanced
dairy cows’ behavioral readiness to access milking com-
partments of an automated milking system (Uetake,
Hurnik, & Johnson, 1997). The social activity of labora-
tory mice, attraction to other mice, huddling, and sexual
behavior were all significantly enhanced by classical
music, whereas rock music produced a decrease in sex-
ual activity and an increase in aggressive huddling
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(Peretti & Kippschull, 1991). “Stereo music” (which
included classical, country, ethnic, “oldies,” and soft
music) reduced agitated and aggressive behaviors in cap-
tive chimpanzees (Howell, Schwandt, Fritz, Roeder, &
Nelson, 2003), although the effectiveness of the various
types of music was not formally compared.
Decreased withdrawal, anxious, or aggressive behav-
ior is also consistent with an increase in positive affect, or
preference. For instance, dogs in a rescue shelter
appeared to prefer (spent more time resting, less time
standing and barking) classical music over silence, and
silence over heavy metal music (Wells, Graham, &
Hepper, 2002). The authors speculate that the “dislike”
of heavy metal music (inferred from increased time
spent barking) was due to dogs’ arousal levels being
increased above optimal levels, whereas classical music
decreased their arousal to a calmer state. Adult caged
hens reared with visual and auditory (radio) stimulation
showed decreased fear reactions (Reed, Wilkins, Austin,
& Gregory, 1993), although the visual and auditory com-
ponents were not separated in this study. Physiological
indices of fear or stress were also reduced more in laying
hens by country music than by classical/jazz/no music
(Ladd et al., 1992), whereas Campo, Gil, and Davila
(2005) found that classical music had no effect on the
same stress index (heterophil-lymphocyte ratio). The
heart rate of singly housed baboons exposed to “oldies
type” radio music was significantly lower than in its
absence (Brent & Weaver, 1996), although behavior and
blood pressure were not affected. Monkeys also showed
a decrease in plasma cortisol levels and abnormal behav-
iors when they were able to control presentation of pop-
ular music (Line, Markowitz, Morgan, & Strong, 1991).
The sensitivity to music type apparent in this litera-
ture raises the possibility of an interaction between ani-
mal handler music preference and animal behavior.
That is, perhaps handlers who listen to country or classi-
cal music become more relaxed, and it is the subsequent
change in handler behavior that affects the animals
(Tosi, Mattiello, Ferrante, & Canali, 2003). Alternatively,
the music to which handlers usually listen will also be
commonly heard by their animals; a “preference” may
therefore develop due to prior exposure or familiarity
rather than any innate musical preference. However,
familiarity and prior exposure are also major determi-
nants of musical preferences of human listeners, so this
possibility simply supports the utility of animal models in
this context and indicates that ecological validity is not a
requirement for auditory-induced effects on animals. It
is noteworthy, however, that when given the choice be-
tween ethologically relevant sounds (including silence)
or music, animals will often prefer the former (Newberry,
1995). For instance, chicks exposed to maternal hen
calls, slow music, or fast music prehatch all showed
 Rickard et al. / FACILITATORY EFFECTS OF MUSIC ON COGNITIVE PERFORMANCE
a preference for maternal hen calls posthatch (Jain,
Sharma, & Wadhwa, 2004).
It cannot be concluded with certainty that the
changes in approach or avoidance behaviors described
above actually reflect a positive affective response to
music. In addition, many of these studies are subject to
the same criticisms of human studies reviewed in the first
section of this article, particularly with regard to the
absence of suitable auditory control groups and the use
of small sample sizes. Nevertheless, there is a broad
range of animal behaviors and physiological indices that
are influenced by music that overlap in animals and
humans, particularly with regard to perception of musi-
cal features, such as rhythm and timbre. At the very least,
music has been shown to influence animals’ behaviors in
ways that are relevant to their well-being. Given the
uncertainty that still surrounds the nature of emotion in
animals in general, this is perhaps the most we can ex-
pect to conclude at this time.
Effect of Music on Cognitive
Performance in Animals
The use of animals to model aspects of human perfor-
mance is commonplace in neuroscience, and animals
have been used extensively in studies on auditory per-
ception and cognition. However, animal models have
been rarely utilized to investigate the effect of music on
cognitive performance. The limited literature is re-
viewed below.
Spatial Performance
One area in which the utility of animal studies has
been insightful is the exploration of the so-called Mozart
effect phenomenon (see “Spatial Per formance”
section). In fact, it is possible that the debate regarding
the mechanisms underlying the effect may ultimately be
answered via animal research. The major competitor to
the arousal-mood hypothesis is the neurophysiological
priming effect, whereby cortical neuronal networks acti-
vated during spatial-temporal reasoning are proposed to
be primed by exposure to complex music (Leng & Shaw,
1991; Rauscher et al., 1995; see “Neuronal Network
Priming” section). Demonstration of a Mozart effect in a
nonhuman species would provide some support for the
neurophysiological priming hypothesis, in that it is gen-
erally regarded that animals are less likely to experience
changes in arousal as a result of music-related mood or
preference. Although there are grounds on which to
appeal this logic (see “Affective Responses to Music”
section), there remains a clear difference in the degree
to which animals and humans are likely to experience
more complex, cognitively mediated forms of emotional
processing in response to music exposure. This is at least
partly because music is not ethologically relevant to ani-
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249
mals. Moreover, personal and cultural associations with
music are unlikely to occur in animals, and as long as
they have not been exposed to human music prior to the
experiment, animals are unlikely to develop preferences
for certain types of music. Animal models therefore pro-
vide an exciting and novel approach to explore the
Mozart effect, as well as other findings of music-induced
cognitive enhancement.
Carlson, Rama, Artchakov, and Linnankoski (1997)
trained 8 macaques on a delayed response task that mea-
sured working memory. They were then exposed to
silence, white noise, a simple rhythm (repetitions of a 5 s
epoch) or Mozart’s Piano Concerto No 21 in C. The audi-
tory treatment was delivered either 15 min before, or
during, the test session (although the duration is not
reported). There was no effect of any auditory treatment
given pretest. However, when given during the test ses-
sion, music by Mozart significantly impaired perfor-
mance, whereas white noise significantly enhanced per-
formance, compared with silence controls. The authors
argued that these effects are likely to be due to distrac-
tion caused by the complexity of the Mozart piece. In
contrast, they argued that white noise may mask disrup-
tive noises in the laboratory. The simple rhythm treat-
ment had no effect on performance even when given
during the test session, possibly because it was approxi-
mately 6 dB SPL quieter than the other auditory
treatments.
Inasmuch as the effect was not replicated in a nonhu-
man species, it could be argued that the Mozart effect is
mediated by exclusively human characteristics, such as
enhanced mood, rather than by direct neurophysiologi-
cal priming, which should also influence animal perfor-
mance. Although this study is a good illustration of the
utility of animal models in exploring the effects of music
on cognitive performance, the methodology used by
Carlson et al. (1997) deviated significantly from the rec-
ommended methodology (Rauscher, 1998). In particu-
lar, a working memory task was used despite Rauscher’s
(1998) emphasis that the effect appeared to be specific
to spatial-temporal tasks. A different piece of music was
used in the human studies, and the sample size was very
small.
These limitations were partially addressed by
Rauscher, Robinson, and Jens (1998) in a study utilizing
Long-Evans rats. They used the same piece of music as
in the original human studies and measured spatial-
temporal task performance (a T-maze). Rats (30 per con-
dition) were exposed to silence, white noise, a minimal-
ist piece of music (8:24 of a piece by composer Philip
Glass) or Mozart’s sonata (K448; duration 8:24). The
auditory treatments were administered in utero as well as
60 days postpartum for 12 hr per day (during their active
phase) at 65 to 70 dB (although background noise levels
250 BEHAVIORAL AND COGNITIVE NEUROSCIENCE REVIEWS
100
75
Intensity (dB SPL)
50
chick
25
rat
human
0 assigned to conditions, litters were kept together, thus
-25
10.0 100.0 1000.0 10000.0
Frequency (Hz)
Figure 1: Sample Audiograms for Domestic Chick (Fay, 1988), Rat
(Heffner, Heffner, Contos, & Ott, 1994), and Human
(Jackson, Heffner, & Heffner, 1999).
NOTE: The most sensitive frequency range for each species is shown by
the trough of the graph, where sounds are audible at lowest stimulus
intensities.
were not reported, so it is not clear how salient the sound
was to the animals). Testing occurred over 5 consecutive
days, with three trials per day, and music was presented
both prior to and during test sessions. Performance of
rats was not affected by the type of music presented dur-
ing the task (i.e., while in the maze), indicating that they
were not distracted by it as presumed for the monkeys in
the study by Carlson et al. (1997). In contrast, the rats
exposed to the Mozart sonata prior to the task com-
pleted the maze faster, and made fewer errors, than did
rats exposed to either minimalist music or white noise
prior to the task. In addition, rats exposed to silence per-
formed similarly to those exposed to white noise, which
excludes the possibility that the difference between the
white noise (or minimalist music) and Mozart-exposed
rats was due to the former performing worse (e.g., due to
either stress or overly low arousal levels). These findings
suggest that the Mozart effect generalizes to an animal
species, which argues against the mechanism being
solely due to musical preference or mood. In particular,
the authors interpret the findings as consistent with stud-
ies that have shown increased hippocampal plasticity
due to environmental enrichment. In addition, they ini-
tially set their study within the same neurophysiological
framework as their human studies (Rauscher et al.,
1995); that is, that music strengthens neural networks in
the temporal lobe, which are also required for spatial
performance. It would be valuable, however, to test ani-
mals’ preference to the different auditory treatments, or
to measure physiological arousal levels, to more con-
clusively reject the mood-arousal hypothesis.
A number of concerns have been raised about the
interpretation of this study (Steele, 2003). First, the rat’s
perceptual frequency range is significantly higher than
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that of humans (see Figure 1). Steele notes that the hear-
ing threshold at 65 to 70 dB reported in Rauscher’s study
for adult Long-Evans (hooded) rats would be around
500 Hz. Therefore, the lower four octaves of the piano
keyboard (which contribute to more than two thirds of
the notes in the piano piece used) would not be heard.
Furthermore, rats are born deaf and therefore would
not have benefited from the in utero exposure to music.
Second, despite claims that animals were randomly
auditory exposure was confounded with litter effects.
Third, the duration of auditory exposure (12 hr per day
in utero and 65 days postpartum) was considerably
greater than that used in the human studies (8:24 once
prior to training). That such chronic exposure during
development is analogous to acute single exposure in a
mature adult is unlikely. The rat data may in fact be more
analogous to long-term music education programs in
children, for which there is also some positive (though
largely correlational) evidence of cognitive-enhancing
effects in humans (see “Music Training” section).
In an attempt to address the limitations of previous
animal studies into the Mozart effect, Macdonald and
Rickard (2005) performed a series of studies with the
domestic chick. Audiograms reveal that, unlike rats, neo-
nate chicks are able to hear frequencies between 100 and
1,000 Hz at moderate intensities (see Figure 1). Chicks
would therefore be able to perceive the majority (at least
90%) of the Mozart sonata when played at intensities
above 55 dB, although the lower notes (i.e., below G#2)
may have been inaudible.
Performance on both spatial-temporal (a simple
maze) and passive avoidance learning was tested to de-
termine if any effect was generalized to nonspatial per-
formance. Macdonald and Rickard (2005) exposed
chicks to either silence, minimalist music, or the Mozart
sonata immediately prior to the training session, or
immediately prior to the testing session. To avoid any
potential “distraction” effects, music was not played dur-
ing testing sessions on either task. The duration of the
auditory treatments was reduced to 1 min, because pre-
vious studies had shown that retention of chicks was
enhanced by auditory treatments lasting less than 2 min
(Toukhsati & Rickard, 2001) and it is important to
ensure the presentation of the auditory stimuli is species-
appropriate (Steele, 2003). Neither pretraining nor pre-
test exposure to the Mozart sonata or minimalist music
enhanced either spatial or passive avoidance perfor-
mance of young chicks (Macdonald & Rickard, 2005).
This finding would seem to lend support to the arousal-
mood theory of the Mozart effect literature in humans,
as no direct neurophysiological priming effect was ob-
served. In addition, it persuasively demonstrates the util-
ity of animal models in music neuroscience research.
 Rickard et al. / FACILITATORY EFFECTS OF MUSIC ON COGNITIVE PERFORMANCE 251

Learning and Memory 1.0

A *
One other area in which animal models have pro-

Mean Discrimination Ratio

0.9
vided insight is in relation to the effect of musical stimuli
on learning and memory. Bates and Horvath (1971)
0.8
demonstrated that rats’ performance on a visual discrim-
ination task was impaired by a nonrhythmic chamber
0.7
symphony piece by Schoenberg. The impairment fol-
lowed presentation of either a melodic or monotonic
0.6
version of this piece, suggesting that rhythmicity was
more critical to rats’ performance than its melodic con-
0.5
tour. In support, exposure to a (rhythmic) Mozart sym- C-R I-R W-N H-E-N Control
T ype of auditory stimulus
phony had no deleterious effect on performance, 1.0

regardless of whether it was presented in melodic or B *

Administration
Time
-1m
monotonal form. * +5 m

Mean Discrimination Ratio

0.9

Schreckenberg and Bird (1987) found that exposing

mice to nonrhythmic music also produced impair- 0.8

ments on a spatial learning task. Music that was “non-

synchronized” (that is, consisting of tones that were not 0.7

aligned with the dominant beat) produced learning and

memory deficits on a maze task, whereas no such deficits 0.6

were observed following exposure to synchronized

music. However, in both these studies, other manipula- 0.5
Control 0.50 1.00 2.00 3.00 5.00
tions of the music were not explored. For instance, it is 1.0
Duration of stimulus (mins)
not known upon which musical properties (e.g., loud- C * *
ness, accent, tempo, time of administration, duration) * *
Mean Discrimination Ratio

0.9
* * *
the effect was dependent. Furthermore, Cross et al. * *
(1967) attributed rats’ preference for music by Mozart 0.8 *
over that of Schoenberg to the difference in rhythmicity,
but there remains a number of other musical properties 0.7

(e.g., timbre) that differ across pieces. Clearly, there is a

need for potential confounding factors to be excluded 0.6

in animal as well as human studies for conclusions to be

drawn with greater confidence. 0.5
Cntrl. -30 -10 -5 -2 -1 0 1 2 5 10 20 40
By reducing music to component elements (such as Time of presentation (mins after training)

basic rhythms with pure tones), many confounds of the

abovementioned studies can be excluded. In humans,
an examination of the individual musical elements re-
sponsible for memory-enhancing effects is problematic,
as this approach risks producing overly simple and un-
interesting stimuli that may be disliked (Steck &
Machotka, 1975). In contrast, such stimuli—particularly
if melodically simple—may be more appropriately
within animals’ perceptual and preferential range. Ani-
mal models may therefore be ideal for investigating the
impact of varying one music element on learning or
memory while holding all other elements constant.
Using this approach, Toukhsati and Rickard (2001)
presented a range of auditory stimuli to neonate chicks
(1- to 2-day-old) in an attempt to improve memory for a
discriminated passive avoidance task. Although a com-
plex rhythmic stimulus (a 1,920 ms sequence comprising
tones set at 120 and 240 ms intervals) presented 5 min
after training significantly enhanced long-term memory,
an isochronous rhythm, white noise, or background
Figure 2
NOTE: Memory-enhancing effects of a synthesized rhythmic auditory
stimulus in chicks are dependent on type of auditory stimulus, dura-
tion, and time of presentation relative to passive avoidance training
(from Toukhsati & Rickard, 2001). C-R = complex rhythmic stimulus; I-
R = isochronous stimulus; W-N = white noise; H-E-N = human environ-
mental noise; Control = no systematic noise. (*Significantly different,
p < .05).
noise had no such memory-facilitating effects (see Fig-
ure 2A). Systematic investigation into the optimum
memory-enhancing acoustic parameters of the rhythmic
stimulus revealed that retention was best facilitated
when the rhythmic stimulus was presented at intensities
between 5 and 15 dBA above background laboratory
noise, and at a frequency of 1 kHz. The effect was not
moderated by the removal or addition of an accent
(Toukhsati & Rickard, 2004). Enhancement was also
dependent on the duration and presentation time of the
auditory stimulus. Retention was enhanced by exposure

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252 BEHAVIORAL AND COGNITIVE NEUROSCIENCE REVIEWS

Neurobiological Mechanisms
Ethologically relevant
Rhythmic Underlying the Effect of Music on
Non-rhythmic
1.0 Cognitive Performance in Animals
Approach calls
Task-relat ed Research on potential mediators of music’s cognitive-
long-term memory index

0.9 enhancing effect in humans has generated numerous

0.8
0.7 Avoidance
calls
0.6
0.5
NSN ICS Asy CRS Fear PC RC FM
Auditory stimulus
Figure 3: The Effects of Various Auditory Stimuli on Long-Term
Memory for the Passive Avoidance Task in Neonate Chicks.
NOTE: Each stimulus was presented for 1 min within 10 after training.
NSN = no systematic noise control; ICS = isochronous rhythm; Asy =
asynchronous version of the CRS; CRS = complex rhythmic stimulus;
Fear = species-specific fear call; PC = species-specific predator call; RC =
species-specific roosting call; FM = species-specific follow-me call; FC =
species-specific food call; 1 kHz = monotonic (1 kHz) transposition of
FC.
to 30 s or 1 min of the rhythmic stimulus, but not by lon-
ger duration periods (2 to 5 min; see Figure 2B). The
treatment enhanced memory if presented between 10
min before and 20 min after training but was not effec-
tive beyond this time window (Toukhsati & Rickard,
2001; see Figure 2C). The rhythmic element of the stim-
ulus was also manipulated in various ways by modifying
the degree of synchronicity, metricity, and repetitiveness
of the stimulus, and by comparing rhythmic and non-
rhythmic maternal hen calls (Rickard, 2005; Toukhsati &
Rickard, 2005). When viewed together, the most parsi-
monious explanation of the memory-enhancing proper-
ties of certain auditory stimuli is that they tend to be
rhythmic (see Figure 3). The high degree of control and
statistical power possible in this experimental design
appears to be responsible for the clearer, more repli-
cable set of findings than is typical in human studies on
this topic. “Music” could also be designed with species-
specific emotional vocalizations and auditory prefer-
ence in mind. The potential for such auditory stimuli,
which have greater meaning to the animal, to have an
even more powerful effect on cognitive performance is
intriguing (see Panksepp & Bernatzky, 2002). However,
the purpose of this review is to explore the utility of
animal models in examining the effects of human music,
and therefore this question is peripheral to the current
discussion.
testable hypotheses. Although some insight has certainly
been gained into the mechanisms underlying this effect,
animal research provides a highly complementary ap-
proach to this investigation (Hodges, 1996; Panksepp &
Bernatzky, 2002; Weinberger, 1999).
Physiological Arousal and
Neurochemical Mediation
As with humans, arousal plays a major mediating role
FC 1kHz
in the effect of music on task performance in ani-
mals. Increased arousal levels are at least one means by
which treatments can enhance memory, with a range of
arousal-inducing manipulations (e.g., administration
of arousal hormones, isolation stress, retraining trials)
improving consolidation of long-term memory in ani-
mals (Bianchin, Mello e Souza, Medina, & Izquierdo,
1999; Crowe, Ng, & Gibbs, 1990; Kobayashi & Kobayashi,
2001; Sandi & Rose, 1994). These findings are consistent
with human literature in which memory is thought to be
modulated via the actions of certain adrenal stress hor-
mones and neurotransmitters (for reviews, see Cahill &
McGaugh, 1998; Castellano, Cabib, & Puglisi-Allegra,
1996; McGaugh, 1989, 2000), with noradrenaline being
identified as a particularly important memory modula-
tor in both animals and humans (see Gibbs & Summers,
2002; Sara, 1991; Tiplady, Bowness, Stien, & Drummond,
2005).
Evidence that exposure to music can also modify
physiological arousal levels in animals has been accumu-
lating (see “Affective Responses to Music” section). For
instance, Ladd et al. (1992) exposed different groups of
caged white Leghorn laying hens in an environmentally
controlled facility to one of three auditory treatments.
For 8 hours per day, birds heard either country music or
jazz/classical while the control groups received no
music. Compared to the control group, hens exposed
to country music had significantly lower heterophil-
lymphocyte ratios, reflecting significantly lower physio-
logical stress levels. In addition, control hens demon-
strated nearly twice as many grooming behaviors as
either music group. Such behavior in excess is also
thought to reflect stress. More recently, Sutoo and
Akiyama (2004) investigated the impact of music on sys-
tolic blood pressure in hypertensive rats. They found
that 2 hr exposure to Mozart’s Adagio from Divertimento No
7 in D Major (K205) (average 65 dB) produced a signifi-
cant decrease in systolic blood pressure and significantly
increased serum calcium and neostriatal dopamine lev-
els pre- to postmusic intervention. These changes were

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 Rickard et al. / FACILITATORY EFFECTS OF MUSIC ON COGNITIVE PERFORMANCE
not present in animals that were not exposed to the
music. The authors concluded that music may affect ani-
mals’ arousal levels and brain function via a calcium/
calmodulin-dependent dopaminergic mediation.
If music modifies neurotransmitters that are also
known to influence performance, a substantial case can
be made that music can, albeit indirectly, affect cogni-
tion. Avian studies, in particular, have consistently con-
firmed that musical stimuli can alter neurotransmitter
levels, with noradrenaline being specifically identified.
Bernatzky, Panksepp, Rossi, and Narayanan (1997)
exposed 2-day-old chicks to either popular instrumental
music or human speech for 30 min twice daily for 10
days. A control group of chicks were exposed to no sys-
tematic sounds. Whole brain neurochemical levels were
then assessed 1 to 2 days following the treatment and
compared with control group levels. Chicks exposed to
music displayed increased levels of noradrenaline
(400%), the noradrenaline metabolite monohydroxy-
phenylglycol (600%), dopamine (200%), and its metab-
olite, homovanillic acid (200%). In contrast, no sig-
nificant neurochemical changes were seen in chicks
exposed to human speech.
Bernatzky, Panksepp, Burgdorf, Nordholm, and Jung
(1998) extended these findings by investigating whether
the neurochemical effects following exposure to popu-
lar instrumental music could be replicated using classi-
cal music (Mozart’s Kroenungskonzert). Chicks were
exposed to 30 min of either music or human speech
twice a day for 10 days. Although the effects were attenu-
ated, whole brain noradrenaline levels of the chicks
exposed to music were still significantly elevated (31%),
compared to those exposed to human speech. In addi-
tion, Panksepp and Bernatzky (2002) found that expo-
sure to both popular music and classical music (Beetho-
ven’s Hammerklavier) elicited behavioral responses (an
increase in head shakes and yawns) similar to those
evoked by infusions of neurochemical agents, which are
known to reduce separation distress in birds. Although
their hypothesis is yet to be directly tested, Panksepp and
Bernatzky postulate that these behavioral responses may
reflect the modulatory effects of music on the chick
neuropeptide system.
Given the marked differences in chick noradrenergic
levels following exposure to either popular or classical
music in Bernatzky et al.’s experiments (1997, 1998), the
specific contribution of various musical elements such as
pitch, rhythm, tempo, or timbre in this context is also of
interest. Of all the components of music, rhythm is his-
torically the most physiologically arousing (Gaston,
1951) and the best predictor of changes in arousal
(Weigl, 1959). In humans, exposure to highly rhythmic
music has resulted in changes in heart rate (e.g., Gerra
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253
et al., 1998; Landreth & Landreth, 1974), respiration
rate (e.g., Iwanaga & Moroki, 1999), skin conductance
(e.g., Henkin, 1957), and neuroendocrine release (e.g.,
Gerra et al., 1998). We therefore explored whether
noradrenergic activation accounted for the memory-
enhancing effects of rhythmic stimuli previously
observed (see “Learning and Memory” section). Consis-
tent with Bernatzky et al.’s (1997, 1998) findings that
musical stimuli increased whole brain noradrenaline
levels in neonate chicks, a higher dose of the b-
adrenergic receptor antagonist, propranolol, was
required to block memory in the presence of a rhythmic
auditory stimulus than in its absence (Toukhsati,
Rickard, Perini, Ng, & Gibbs, 2005). Furthermore, the
duration of an intermediate phase of memory forma-
tion, typically sensitive to arousal levels, was extended in
the presence of the rhythmic stimulus. The memory-
enhancing effect of a rhythmic maternal hen call was
also prevented by blockade of b-adrenoceptors (Field,
Rickard, Toukhsati, & Gibbs, 2005). Taken together, this
series of experiments provides strong evidence that com-
plex rhythmic auditory stimuli enhance memory by pro-
moting moderate levels of physiological arousal through
m o d u l a t i o n o f no r a d r e n a l i n e ac t i v i t y at b -
adrenoceptors. In addition, these studies highlight the
efficacy of using an animal model to investigate the
effects that exposure to musical stimuli can have on
cognitive performance.
Neural Plasticity
Animal models also provide an opportunity to
explore the speculation that music might enhance cog-
nition independently of the emotion/arousal path-
way. It has, after all, been shown that environmental
enrichment can induce neuronal plasticity in animals
(Greenough, McDonald, Parnisari, & Camel, 1986;
Meaney, Aitken, Bhatnagar, vanBerkel, & Sapolsky, 1988;
Rosenzweig, 1984; Rosenzweig & Bennett, 1996) and
possibly humans (Raine et al., 2001; Schaie, 1994). In
support, Wadhwa and colleagues observed a number of
neuroanatomical changes in the chick brain in response
to auditory stimuli. Prenatal exposure to musical (sitar)
or species-specific stimuli resulted in larger nuclei of
medio-rostral neostriatum/hyperstriatum ventrale
neurons than in the presence of background noise
(Panicker, Wadhwa, & Roy, 2002). This region has been
implicated in learning and memory in the young chick
(Rose & Csillag, 1985; Sedman, O’Dowd, Rickard, Gibbs,
& Ng, 1992). Panicker et al. (2002) also found a greater
number of PV- and CaBP-staining neurons in music-
exposed animals than in control animals; these proteins
act as intracellular buffers, enabling neurons to display
high electrical activity without calcium overload. The
254 BEHAVIORAL AND COGNITIVE NEUROSCIENCE REVIEWS

authors hypothesized that this may enhance potential Summary

for long-term potentiation, which is widely believed to
The material reviewed in this section indicates that
underlie some forms of memory formation.
animal models provide a genuine contribution to under-
Prenatal exposure to either species-specific or musi-
standing the cognitive and physiological effects of musi-
cal auditory stimuli increased the length, number, and
cal stimuli. Although there are important differences in
size of brainstem auditory nuclei, as well as the number
the way that relations between pitches are perceived, ani-
of glia and neuropil, and the normal pattern of synaptic
mals do appear to hear at least the basic features of
protein expression (syntaxin, synaptophysin) in
music, including pitch, loudness, and temporal patterns.
brainstem nuclei (Alladi, Wadhwa, & Singh, 2002;
Differences between human and animal perception may
Wadhwa, Anand, & Bhowmick, 1999). Exposure to audi-
be partly due to lack of exposure, as further training can
tory stimuli can also result in neurophysiological
result in the use of relative pitch in animals, as occurs
changes in synaptic plasticity. Engineer et al. (2004)
with developing humans. Nonetheless, it reflects a major
exposed rats to a standard or a visually and acoustically
difference of which researchers should be aware when
enriched environment for 8 weeks. Auditory stimuli in
using animal models in music neuroscience research.
the enriched condition included wind chimes, bells, sim-
There has been limited demonstration to date of
ple tones of various frequencies, noise bursts, classical
cognitive-enhancing effects of music in animals.
music, and other complex sounds such as rat vocaliza-
Reduced forms of music, such as the monotonic but
tions and rustling leaves. Enrichment was found to
rhythmic stimuli used in our laboratory, have, however,
increase a number of indices of auditory cortex
consistently yielded significant improvements in perfor-
response, including amplitude, sensitivity, and number.
mance on a memory task in birds. The utility of animal
The possibility that the benefits of auditory enrichment
models is most cogently demonstrated in the search for
on synaptic plasticity might translate to humans, or that
potential underlying mechanisms of such effects. A con-
nonauditory cortex might also be modified, has not
sistent conclusion across laboratories is that musical
been explored. This research has exciting implications
stimuli, particularly the temporal components such as
for populations (such as the aged) who experience
rhythm and tempo, are arousing and subsequently mod-
cognitive deficits as a result of gradually declining
ify catecholamine levels. Taken together with the dem-
neuronal efficacy.
onstrated effects of music on synaptic plasticity in avian
Schreckenberg and Bird (1987) explored the effect
brains, there is a substantial case for the potential of
of nonsynchronized and synchronized rhythmic music
music to modify cognitive performance.
on hippocampal pyramidal and glial cell morphology
Music perception in humans is hierarchical, with
and cognitive performance in mice. Animals were
higher levels recruiting more advanced perceptual and
exposed to one type of music or silence from birth until 2
cognitive processes (Swartz et al., 1989; Umemoto,
months of age (80-86 dB). They found that exposure to
1990). The basic levels comprise perception, identifica-
nonsynchronized rhythmic music produced a number
tion, and discrimination of changes in pitch, timbre, and
of significant changes in mouse hippocampi: Neuronal
intensity. The middle levels involve perception of mel-
dendrites exhibited abnormal sprouting and branching,
ody, rhythm, and harmony, and higher still is compre-
astroglia were increased in size and number, and
hension of form and structure. An emotional appre-
increased levels of mRNA were also observed relative to
ciation of music may be the highest level of music
the other two groups. No significant morphological
perception (Umemoto, 1990), although some of the
changes occurred in the hippocampi of mice exposed to
most powerful affective responses to music may actually
synchronized rhythmic music or to silence. Mice ex-
be—in neurobiological terms—quite basic and there-
posed to the nonsynchronized rhythmic music also per-
fore could be shared by animals (Panksepp, 2003;
formed worse on a spatial learning task than did mice
Panksepp & Bernatzky, 2002). There is no doubt that
exposed to synchronized rhythmic music or to silence.
human interaction with music produces other experi-
The authors speculated that abnormal branching and
ences, which are more cognitive in nature and not
excessive glial growth may underlie disturbances of nor-
known to occur in animals (Panksepp & Bernatzky,
mal learning and memory. In this context, it is interest-
2002). Nevertheless, a comparative psychology ap-
ing that exposure to uncontrollable noise (which may be
proach to the perception of music reveals that animals
how animals interpret the irregular music) also impairs
do perceive at least the basic components of music in a
human performance on many tasks (Broadbent, 1979;
similar way as we do. Moreover, animal research has been
Eschenbrenner, 1971). Such findings demonstrate that,
valuable in defining those responses that might be
whether the effect is facilitatory or deleterious to cogni-
unique to humans. As long as music research utilizing
tive performance, auditory stimuli can indeed have a
animal models acknowledges this bottom-up approach,
significant impact on brain plasticity.
there is no reason why generalizable conclusions cannot

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 Rickard et al. / FACILITATORY EFFECTS OF MUSIC ON COGNITIVE PERFORMANCE
be drawn. An interesting aside is that some neuro-
degenerative disorders, such as Alzheimer’s disease,
probably strip humans of the higher levels of music
appreciation also (Aldridge, 1995; Swartz et al., 1989). In
this context, animal models may even offer a unique
insight into human cognitive performance.
Well constructed music is uniquely efficacious in reso-
nating with our basic emotional systems, bringing to life
many affective proclivities that may be encoded as birth-
rights, within ancient neural circuits constructed by our
genes, many of which we share homologously with other
mammals. (Panksepp & Bernatzky, 2002, p. 136)
CONCLUSION
The claim that music can enhance human cognitive
performance is enticing but remains controversial. It
would appear that substantial benefits can result from
several different types of music treatment (passive expo-
sure, music therapy, and music training), particularly in
relation to spatial performance and learning and mem-
ory. However, much of this research can be disputed on
the grounds that inadequate experimental designs were
used. Innovative approaches are required to make sub-
stantial progress on this issue, and this review has dem-
onstrated that animal research provides a valuable con-
junct to human research in this particular field. The
influence of auditory stimuli on cognitive performance
in animals is free of many of the human factors that con-
found the interpretation of experimental results and
overlay our more primal responses to music. This
approach is necessarily reductionist, in that it is the holis-
tic and interactionist nature of music and the listener
that confounds much of the human research. Compara-
tive studies have demonstrated that this approach is,
however, neither trivial nor absurd. Several programs
utilizing animal models have already illustrated the
enhanced clarity and reproduction that may be possible
with tighter experimental control.
Despite the limited evidence of cognitive-enhancing
effects of music, the potential for music to impact cogni-
tive performance is enormous. Empirical investigations
in both humans and animals converge in that modula-
tion of arousal has been offered as a likely explanation
for a facilitative effect of music on performance.
Whether due to its rhythmic properties or its capacity to
elicit strong emotions, music can produce autonomic
and neurochemical responses that are consistent with an
aroused state, which can in turn result in enhanced task
performance. Music exposure, and particularly longer
term training, can also modify neuronal efficacy, which
may benefit performance in other domains. The neuro-
biological hypotheses regarding the mechanisms by
which music can improve performance do not, there-
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255
fore, seem to be lacking. This avenue of research
requires extension such that the fundamental and more
complex features of music treatments responsible for
cognitive enhancement are identified. This endeavor
promises to shed light on the components of music that
are critical for particular purposes (e.g., music therapy,
commercial selling, background relaxing music), and
conclusions are likely to be drawn from both human and
animal domains.
Although clearly a complete understanding of the
effects of music cannot be obtained by observing animal
responses, knowledge of the more fundamental or
“direct” effects of music may be enhanced by including
a bottom-up approach. Once these truly universal
responses to music are better defined, a firmer founda-
tion will be available, upon which the more complex
effects of music on humans can be investigated. Empiri-
cal research can be guided by having separated those
generic features of music that produce cognitive, affec-
tive, and physiological responses from those that are
dependent on more complex constructs that vary from
person to person. These more complex layers can,
clearly, only be studied in humans.
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