Why Would Musical Training Benefit The Neural Encoding of Speech? The OPERA Hypothesis

Hypothesis and Theory Article
published: 29 June 2011

doi: 10.3389/fpsyg.2011.00142
Why would musical training benefit the neural encoding of

speech? The OPERA hypothesis
Aniruddh D. Patel*
Department of Theoretical Neurobiology, The Neurosciences Institute, San Diego, CA, USA
Edited by: Mounting evidence suggests that musical training benefits the neural encoding of speech. This
Lutz Jäncke, University of Zurich,
Switzerland
paper offers a hypothesis specifying why such benefits occur.The “OPERA” hypothesis proposes
that such benefits are driven by adaptive plasticity in speech-processing networks, and that
Reviewed by:
Kai Alter, Newcastle University, UK this plasticity occurs when five conditions are met. These are: (1) Overlap: there is anatomical
Stefan Koelsch, Freie Universität Berlin, overlap in the brain networks that process an acoustic feature used in both music and speech
Germany (e.g., waveform periodicity, amplitude envelope), (2) Precision: music places higher demands on
*Correspondence: these shared networks than does speech, in terms of the precision of processing, (3) Emotion:
Aniruddh D. Patel, Department of
the musical activities that engage this network elicit strong positive emotion, (4) Repetition:
Theoretical Neurobiology, The
Neurosciences Institute, 10640 John the musical activities that engage this network are frequently repeated, and (5) Attention: the
Jay Hopkins Drive, San Diego, CA musical activities that engage this network are associated with focused attention. According
92121, USA. to the OPERA hypothesis, when these conditions are met neural plasticity drives the networks
e-mail: [email protected]
in question to function with higher precision than needed for ordinary speech communication.
Yet since speech shares these networks with music, speech processing benefits. The OPERA
hypothesis is used to account for the observed superior subcortical encoding of speech in
musically trained individuals, and to suggest mechanisms by which musical training might
improve linguistic reading abilities.
Keywords: music, speech, neural plasticity, neural encoding, hypothesis
Introduction can be enhanced by non-linguistic auditory training (i.e., learning

Recent EEG research on human auditory processing suggests that to play a musical instrument or sing). This has practical significance
musical training benefits the neural encoding of speech. For exam- because the quality of brainstem speech encoding has been directly
ple, across several studies Kraus and colleagues have shown that the associated with important language skills such as hearing in noise
neural encoding of spoken syllables in the auditory brainstem is and reading ability (Banai et al., 2009; Parbery-Clark et al., 2009).
superior in musically trained individuals (for a recent overview, see This suggests that musical training can influence the development
Kraus and Chandrasekaran, 2010; this is further discussed in The of these skills in normal individuals (Moreno et al., 2009; cf. Strait
Auditory Brainstem Response to Speech: Origins and Plasticity). et al., 2010; Parbery-Clark et al., 2011). Furthermore, as noted by
Kraus and colleagues have argued that experience-dependent neu- Kraus and Chandrasekaran (2010), musical training appears to
ral plasticity in the brainstem is one cause of this enhancement, strengthen the same neural processes that are impaired in individu-
based on the repeated finding that the degree of enhancement als with certain speech and language processing problems, such as
correlates significantly with the amount of musical training (e.g., developmental dyslexia or hearing in noise. This has clear clinical
Musacchia et al., 2007, 2008; Wong et al., 2007; Strait et al., 2009). implications for the use of music as a tool in language remediation.
They suggest that plasticity in subcortical circuits could be driven Kraus and colleagues have provided a clear hypothesis for how
by descending (“corticofugal”) neural projections from cortex onto musical training might influence the neural encoding of speech
these circuits. There are many such projections in the auditory (i.e., via plasticity driven by corticofugal projections). Yet, from
system (exceeding the number of ascending fibers), providing a a neurobiological perspective, why would musical training drive
potential pathway for cortical signals to tune subcortical circuits adaptive plasticity in speech processing networks in the first place?
(Winer, 2006; Kral and Eggermont, 2007; Figure 1). Kraus and Chandrasekaran (2010) point out that both music and
The arguments of Kraus and colleagues have both theoretical speech use pitch, timing, and timbre to convey information, and
and practical significance. From a theoretical standpoint, their pro- suggest that years of processing these cues in a fine-grained way
posal contradicts the view that auditory processing is strictly hierar- in music may enhance their processing in the context of speech.
chical, with hardwired subcortical circuits conveying neural signals The current “OPERA” hypothesis builds on this idea and makes it
to cortical regions in a purely feed-forward fashion. Rather, their more specific. It proposes that music-driven adaptive plasticity in
ideas support a view of auditory processing involving rich two-way speech processing networks occurs because five essential condi-
interactions between subcortical and cortical regions, with struc- tions are met. These are: (1) Overlap: there is overlap in the brain
tural malleability at both levels (cf. the “reverse hierarchy theory” networks that process an acoustic feature used in both speech and
of auditory processing, Ahissar et al., 2009). From a practical stand- music, (2) Precision: music places higher demands on these net-
point, their proposal suggests that the neural encoding of speech works than does speech, in terms of the precision of processing,
www.frontiersin.org June 2011 | Volume 2 | Article 142 | 1

Patel Musical training and speech encoding
Prior to these core sections of the paper, section “The Auditory

Brainstem Response to Speech: Origins and Plasticity” provides
background on the auditory brainstem response to speech and evi-
dence for neural plasticity in this response. Following the two core
sections, section “Musical vs. Linguistic Training for Speech Sound
Encoding” addresses the question of the relative merits of musical
vs. linguistic training for improving the neural encoding of speech.
The auditory brainstem response to speech: origins

and plasticity
As noted by Chandrasekaran and Kraus (2010), “before speech can
be perceived and integrated with long-term stored linguistic rep-
resentations, relevant acoustic cues must be represented through
a neural code and delivered to the auditory cortex with temporal
and spectral precision by subcortical structures.” The auditory
pathway is notable for complexity of its subcortical processing,
which involves many distinct regions and a rich network of ascend-
ing and descending connections (Figure 2). Using scalp-recorded
EEG, population-level neural responses to speech in subcortical
regions can be recorded non-invasively from human listeners (for
a tutorial, see Skoe and Kraus, 2010). The brainstem response to
the spoken syllable /da/ is shown in Figure 3, and the details of
this response near syllable onset are shown in Figure 4. (Note that
the responses depicted in these figures were obtained by averag-
ing together the neural responses to thousands of repetitions of
the syllable /da/, in order to boost the signal-to-noise ratio in the
scalp-recorded EEG).
Broadly speaking, the response can be divided into two com-
Figure 1 | A simplified schematic of the ascending auditory pathway
between the cochlea and primary auditory cortex, showing a few of the
ponents: a transient onset response (commencing 5–10 ms after
subcortical structures involved in auditory processing, such as the syllable onset, due to neural delays) and an ongoing component
cochlear nuclei in the brainstem and the inferior colliculus in the known as the frequency-following response (FFR). The transient
midbrain. Solid red lines show ascending auditory pathways, dashed lines onset response reflects the synchronized impulse response of neu-
show descending (“corticofugal”) auditory pathways. (In this diagram,
ronal populations in a number of structures, including the coch-
corticofugal pathways are only shown on one side of the brain for simplicity.
See Figure 2 for a more detailed network diagram). From Patel and Iversen
lea, brainstem, and midbrain. This response shows electrical peaks
(2007), reproduced with permission. which are sensitive to the release burst of the /d/ and the formant
transition into the vowel. The FFR is the summed responses of elec-
trical currents from structures including cochlear nucleus, supe-
(3) Emotion: the musical activities that engage this network elicit rior olivary complex, lateral lemniscus, and the inferior colliculus
strong positive emotion, (4) Repetition: the musical activities that (Chandrasekaran and Kraus, 2010). The periodicities in the FFR
engage this network are frequently repeated, and (5) Attention: reflect the synchronized component of this population response,
the musical activities that engage this network are associated with arising from neural phase locking to speech waveform periodicities
focused attention. According to the OPERA hypothesis, when these in the auditory nerve and propagating up the auditory pathway
conditions are met neural plasticity drives the networks in question (cf. Cariani and Delgutte, 1996). The form of the FFR reflects the
to function with higher precision than needed for ordinary speech transition period between burst onset and vowel, and the vowel
communication. Yet, since speech shares these networks with music, itself, via reflection of the fundamental frequency (F0) and some
speech processing benefits. of the lower-frequency harmonics of the vowel.
The primary goals of this paper are to explain the OPERA The auditory brainstem response occurs with high reliability
hypothesis in detail (section The Conditions of the OPERA in all hearing subjects (Russo et al., 2004), and can be recorded in
Hypothesis), and to show how it can be used to generate predictions passive listening tasks (e.g., while the listener is watching a silent
to drive new research (section Putting OPERA to Work: Musical movie or even dozing). Hence it is thought to reflect sensory, rather
Training and Linguistic Reading Skills). The example chosen to than cognitive processing (though it may be shaped over longer
illustrate the prediction-generating aspect of OPERA concerns rela- periods of time via cognitive processing of speech or music, as
tions between musical training and linguistic reading skills (cf. discussed below). Since the response resembles the acoustic signal
Goswami, 2010). One motivation for this choice is to show that in several respects (e.g., in its temporal and spectral structure),
the OPERA hypothesis, while developed on the basis of research on correlation measures can be used to quantify the similarity of the
subcortical auditory processing, can also be applied to the cortical neural response to the spoken sound, and hence to quantify the
processing of acoustic features of speech. quality of speech sound encoding by the brain (Russo et al., 2004).
Frontiers in Psychology | Auditory Cognitive Neuroscience June 2011 | Volume 2 | Article 142 | 2

Figure 2 | Schematic diagram of the auditory system, illustrating the

many subcortical processing stations between the cochlea (bottom) and
cortex (top). Blue arrows represent ascending (bottom-up) pathways; red
arrows represent descending projections. From Chandrasekaran and Kraus
(2010), reproduced with permission.
As noted previously, the quality of this encoding is correlated with

important real-world language abilities, such as hearing in noise
and reading. For example, Banai et al. (2009) found that the latency
of specific electrical peaks in the auditory brainstem response to
/da/ predicted word reading scores, when the effects of age, IQ, and
the timing of click-evoked brainstem responses were controlled via
partial correlation.
Figure 3 | The auditory brainstem response to the spoken syllable /da/
Several recent studies have found that the quality of subcortical (red) in comparison to the acoustic waveform of the syllable (black). The
speech sound encoding is significantly greater in musically trained neural response can be studied in the time domain as changes in amplitude
individuals (e.g., Musacchia et al., 2007; Wong et al., 2007; Lee et al., across time (top, middle and bottom-left panels) and in the spectral domain as
2009; Parbery-Clark et al., 2009; Strait et al., 2009). To take one spectral amplitudes across frequency (bottom-right panel). The auditory
brainstem response reflects acoustic landmarks in the speech signal with
example, Parbery-Clark et al. (2009) examined the auditory brain-
submillisecond precision in timing and phase locking that corresponds to (and
stem response in young adults to the syllable /da/ in quiet and in physically resembles) pitch and timbre information in the stimulus. From Kraus
background babble noise. When listening to “da” in noise, musi- and Chandrasekaran (2010), reproduced with permission.
cally trained listeners showed shorter-latency brainstem responses
to syllable onset and formant transitions relative to their musically
untrained counterparts, as well as enhanced representation of speech encoding in this study, and the study of Parbery-Clark et al. (2009),
harmonics and less degraded response morphology. The authors sug- were evident in the absence of attention to the speech sounds or any
gest that these differences indicate that musicians had more synchro- overt behavioral task.
nous neural responses to the syllable. To take another example, Wong One interpretation of the above findings is that they are entirely due
et al. (2007) examined the FFR in musically trained and untrained to innate anatomical or physiological differences in the auditory systems
individuals. Prior work had shown that oscillations in the FFR track of musicians and non-musicians. For example, the shorter latencies
the F0 and lower harmonics of the voice dynamically over the course and stronger FFR responses to speech in musicians could be due to
of a single syllable (Krishnan et al., 2005). In the study of Wong et al. increased synchrony among neurons that respond to harmonically
(2007), native English speakers unfamiliar with Mandarin listened complex sounds, and this in turn could reflect genetically mediated
passively to the Mandarin syllable /mi/ (pronounced “me”) with three differences in the number or spatial distribution of synaptic connections
different lexical tones. The salient finding was that the quality of F0 between neurons. Given the heritability of certain aspects of cortical
tracking was superior in musically trained individuals (Figure 5). It is structure (e.g., cortical thickness, which may in turn reflect the amount
worth noting that musician–non-musician differences in brainstem of arborization of dendrites, Panizzon et al., 2009), it is plausible that

Figure 4 | Time–amplitude waveform of a 40-ms synthesized speech time delay to the auditory brainstem. The start of the formant transition period
stimulus /da/ is shown in blue (time shifted by 6 ms to be comparable is marked by wave C, marking the change from the burst to the periodic portion
with the neural response). The first 10 ms of the syllable are characterized by of the syllable, that is, the vowel. Waves D, E, and F represent the periodic
the onset burst of the consonant /d/; the following 30 ms are the formant portion of the syllable (frequency-following response) from which the
transition to the vowel /a/. The time–amplitude waveform of the time-locked fundamental frequency (F0) of the stimulus can be extracted. Finally, wave O
brainstem response to the 40-ms /da/ is shown below the stimulus, in black. marks stimulus offset. From Chandrasekaran and Kraus (2010), reproduced
The onset response (V) begins 6–10 ms following the stimulus, reflecting the with permission.
Figure 5 | Frequency-following responses to the spoken syllable /mi/ with lines) of the FFR’s primary periodicity, from the same two individuals. For the
a “dipping” (tone 3) pitch contour from Mandarin. The top row shows FFR musician, the FFR waveform is more periodic and its periodicity tracks the
waveforms of a musician and non-musician subject; the bottom row shows the time-varying F0 contour of the spoken syllable with greater accuracy. From
fundamental frequency of the voice (thin black line) and the trajectories (yellow Wong et al. (2007), reproduced with permission.

some proportion of the variance in brainstem responses to speech is due of tone languages.) For example, the syllable “pesh” meant glass,
to genetic influence on subcortical neuroanatomy. In other words, it is pencil, or table depending on whether it was spoken with a level,
plausible that some individuals are simply born with sharper hearing rising, or falling–rising tone. Participants underwent a training
than others, which confers an advantage in processing any kind of com- program in which each syllable, spoken with a lexical tone, was
plex sound. If sharper hearing makes an individual more likely to pursue heard while viewing a picture of the word’s meaning. Quizzes
musical training, this could explain the positive association between were given periodically to measure word learning. The FFR to an
musical training and the superior neural encoding of speech sounds, untrained Mandarin syllable (/mi/) with tones 1–3 was measured
in the absence of any experience-dependent brainstem plasticity. before and after training. The salient finding was that the quality
Alternatively, musical training could be a cause of the enhanced of FFR tracking of fundamental frequency (F0) of Mandarin tones
neural encoding of speech sounds. That is, experience-dependent improved with training. Compared to pretraining measures, par-
neural plasticity due to musical training might cause changes in ticipants showed increased energy at the fundamental frequency
brain structure and/or function that result in enhanced subcortical of the voice and fewer F0 tracking errors (Figure 7). Song et al.
speech processing. There are several reasons to entertain this pos- (2008) note that this enhancement likely reflects enhanced syn-
sibility. First, there is recent evidence that the brainstem encoding chronization of neural firing to stimulus F0, which could in turn
of speech sounds shows training-related neural plasticity, even in result from more neurons firing at the F0 rate, more synchronous
adult listeners. This was demonstrated by Song et al. (2008), who firing, or some combination of these two. A striking aspect of these
had native English speakers learn a vocabulary of six nonsense findings is the small amount of training involved in the study: just
syllables, each paired with three lexical tones based on Mandarin eight 30-min sessions spread across 2 weeks. These results show
tones 1–3 (Figure 6). (The participants had no prior knowledge that the adult auditory brainstem response to speech is surprisingly
malleable, exhibiting plasticity following relatively brief training.
The second reason to consider a role for musical training in
enhanced brainstem responses to speech is that several studies
which report superior brainstem encoding of speech in musicians
also report that the degree of enhancement correlates significantly
with the amount of musical training (e.g., Musacchia et al., 2007,
2008; Wong et al., 2007; Lee et al., 2009; Strait et al., 2009). Finally,
the third reason is that longitudinal brain-imaging studies have
shown that musical training causes changes in auditory corti-
cal structure and function, which are correlated with increased
auditory acuity (Hyde et al., 2009; Moreno et al., 2009). As noted
previously there exist extensive corticofugal (top-down) projec-
tions from cortex to all subcortical auditory structures. Evidence
from animal studies suggests that activity in such projections can
tune the response patterns of subcortical circuits to sound (see
Tzounopoulos and Kraus, 2009 for a review, cf. Kral and Eggermont,
2007; Suga, 2008; Bajo et al., 2010).
Figure 6 | F0 contours of the three Mandarin tones used in the study of Hence the idea that musical training benefits the neural encod-
Song et al. (2008): high-level (tone 1), rising (tone 2), and dipping/falling–rising
ing of speech is neurobiologically plausible, though longitudinal
(tone 3). Reproduced with permission.
randomized controlled studies are needed to establish this with
Figure 7 | Brainstem encoding of the fundamental frequency (F0) the panel), the post-training FFR in the same participant (right panel) shows
Mandarin syllable /mi/ with a “dipping” (tone 3) pitch contour. The F0 more faithful tracking of time-varying F0 contour of the syllable. Data from a
contour of the syllable is shown by the thin black line, and the trajectory of representative participant from Song et al. (2008). Reproduced with
FFR periodicity is shown by the yellow line. Relative to pretraining (left permission.

certainty. To date, randomized controlled studies of the influence periodicity is an important feature of many spoken and musical
of musical training on auditory processing have focused on cortical sounds, and contributes to the perceptual attribute of pitch in
rather than subcortical signals (e.g., Moreno et al., 2009). Hopefully both domains. At the subcortical level, the auditory system likely
such work will soon be extended to include subcortical measures. uses similar mechanisms and networks for encoding periodicity
Many key questions remain to be addressed, such as how much in speech and music, including patterns of action potential timing
musical training is needed before one sees benefits in speech encod- in neurons in the auditory pathway between cochlea and inferior
ing, how large and long-lasting such benefits are, whether cortical colliculus (Cariani and Delgutte, 1996; cf. Figure 2). As noted in
changes precede (and cause) subcortical changes, and the relative section “The Auditory Brainstem Response to Speech: Origins and
effect of childhood vs. adult training. Even at this early stage, how- Plasticity,” the synchronous aspect of such patterns may underlie
ever, it is worth considering why musical training would benefit the FFR.
the neural encoding of speech. Of course, once the pitch of a sound is determined (via mecha-
nisms in subcortical structures and perhaps in early auditory corti-
The conditions of the OPERA hypothesis cal regions, cf. Bendor and Wang, 2005), then it may be processed
The OPERA hypothesis aims to explain why musical training in different ways depending on whether it occurs in a linguistic
would lead to adaptive plasticity in speech-processing networks. or musical context. For example, neuroimaging of human pitch
According to this hypothesis, such plasticity is engaged because perception reveals that when pitch makes lexical distinctions
five essential conditions are met by music processing. These are: between words (as in tone languages), pitch processing shows a
overlap, precision, emotion, repetition, and attention (as detailed left hemisphere bias, in contrast to the typical right-hemisphere
below). It is important to note that music processing does not dominance for pitch processing (Zatorre and Gandour, 2008). The
automatically meet these conditions. Rather, the key point is that normal right-hemisphere dominance in the cortical analysis of
music processing has the potential to meet these conditions, and pitch may reflect enhanced spectral resolution in right-hemisphere
that by specifying these conditions, OPERA opens itself to empirical auditory circuits (Zatorre et al., 2002), whereas left hemisphere
testing. Specifically, musical activities can be designed which fulfill activations in lexical tone processing may reflect the need to inter-
these conditions, with the clear prediction that they will lead to face pitch information with semantic and/or syntactic information
enhanced neural encoding of speech. Conversely, OPERA predicts in language.
that musical activities not meeting the five conditions will not lead The larger point is that perceptual attributes (such as pitch)
to such enhancements. should be conceptually distinguished from acoustic features (such
Note that OPERA is agnostic about the particular cellular as periodicity). The cognitive processing of a perceptual attrib-
mechanisms involved in adaptive neural plasticity. A variety of ute (such as pitch) can be quite different in speech and music,
changes in subcortical circuits could enhance the neural encoding reflecting the different patterns and functions the attribute has
of speech, including changes in the number and spatial distribution in the two domains. Hence some divergence in the cortical pro-
of synaptic connections, and/or changes in synaptic e fficacy, which cessing of that attribute is to be expected, based on whether it is
in turn can be realized by a broad range of changes in synaptic embedded in speech or music (e.g., Peretz and Coltheart, 2003).
physiology (Edelman and Gally, 2001; Schnupp et al., 2011, Ch. On the other hand, the basic encoding of acoustic features under-
7). OPERA makes no claims about precisely which changes are lying that attribute (e.g., periodicity, in the case of pitch) may
involved, or precisely how corticofugal projections are involved involve largely overlapping subcortical circuits. After all, compared
in such changes. These are important questions, but how adaptive to auditory cortex, with its vast numbers of neurons and many
plasticity is manifested in subcortical networks is a distinct question functionally specialized subregions (Kaas and Hackett, 2000),
from why such plasticity is engaged in the first place. The OPERA subcortical structures have far fewer neurons, areas, and connec-
hypothesis addresses the latter question, and is compatible with a tions and hence less opportunity to partition speech and music
range of specific physiological mechanisms for neural plasticity. processing into different circuits. Hence the idea that the sensory
The remainder of this section lists the conditions that must encoding of periodicity (or other basic acoustic features shared
be met for musical training to drive adaptive plasticity in speech by speech and music) uses overlapping subcortical networks is
processing networks, according to the OPERA hypothesis. For the neurobiologically plausible.
sake of illustration, the focus on one particular acoustic feature
shared by speech and music (periodicity), but the logic of OPERA Precision
applies to any acoustic feature important for both speech and music, Let us assume that OPERA condition 1 is satisfied, that is, that
including spectral structure, amplitude envelope, and the timing a shared acoustic feature in speech and music is processed by
of successive events. (In section “Putting OPERA to Work: Musical overlapping brain networks. OPERA holds that for musical
Training and Linguistic Reading Skills,” where OPERA is used to training to influence the neural encoding of speech, music must
make predictions about how musical training might benefit reading place higher demands on the nervous system than speech does,
skills, the focus will be on amplitude envelope). in terms of the precision with the feature must be encoded for
adequate communication to occur. This statement immediately
Overlap raises the question of what constitutes “adequate communication”
For musical training to influence the neural encoding of speech, an in speech and music. For the purposes of this paper, adequate
acoustic feature important for both speech and music perception communication for speech is defined as conveying the semantic
must be processed by overlapping brain networks. For example, and propositional content of spoken utterances, while for music,

it is defined as c onveying the structure of musical sequences. their frequencies and rates of change; Stevens, 1998). While any
Given these definitions, how can one decide whether music places individual cue may be ambiguous, when cues are integrated and
higher demands on the nervous system than speech, in terms interpreted in light of the current phonetic context, they provide a
of the precision of encoding of a given acoustic feature? One strong pointer to the intended phonological category being com-
way to address this question is to ask to what extent a perceiver municated (Toscano and McMurray, 2010; Toscano et al., 2010).
requires detailed information about the patterning of that feature Furthermore, when words are heard in sentence context, listeners
in order for adequate communication to occur. Consider the fea- benefit from multiple knowledge sources (including semantics,
ture of periodicity, which contributes to the perceptual attribute syntax, and pragmatics) which provide mutually interacting con-
of pitch in both speech and music. In musical melodies, the notes straints that help a listener identify the words in the speech stream
of melodies tend to be separated by small intervals (e.g., one or (Mattys et al., 2005).
two semitones, i.e., approximately 6 or 12% changes in pitch, cf. Hence one can hypothesize that the use of context-based cue
Vos and Troost, 1989), and a pitch movement of just one semitone integration and multiple knowledge sources in word recognition
(∼6%) can be structurally very important, for example, when it helps relax the need for a high degree of precision in acoustic
leads to a salient, out-of-key note that increases the complexity of analysis of the speech signal, at least in terms of what is needed
the melody (e.g., a C# note in the key of C, cf. Eerola et al., 2006). for adequate communication (cf. Ferreira and Patson, 2007). Of
Hence for music perception, detailed information about pitch is course, this is not to say that fine phonetic details are not relevant
important. Indeed, genetically based deficits in fine-grained pitch for linguistic communication. There is ample evidence that lis-
processing are thought to be one of the important underlying teners are sensitive to such details when they are available (e.g.,
causes of musical tone deafness or “congenital amusia” (Peretz McMurray et al., 2008; Holt and Idemaru, 2011). Furthermore,
et al., 2007; Liu et al., 2010). these details help convey rich “indexical” information about speaker
How crucial is detailed information about spoken pitch patterns identity, attitude, emotional state, and so forth. Nevertheless, the
to the perception of speech? In speech, pitch has a variety of linguis- question at hand is what demands are placed on the nervous sys-
tic functions, including marking emphasis and phrase boundaries, tem of a perceiver in terms of the precision of acoustic encoding
and in tone languages, making lexical distinctions between words. for basic semantic/propositional communication to occur (i.e.,
Hence there is no doubt that pitch conveys significant informa- “adequate” linguistic communication). How do these demands
tion in language. Yet the crucial question is: to what extent does compare to the demands placed on a perceiver for adequate musi-
a perceiver require detailed information about pitch patterns for cal communication?
adequate communication to occur? One way to address this ques- As defined above, adequate musical communication involves
tion is to manipulate the pitch contour of natural sentences and see conveying the structure of musical sequences. Conveying the struc-
how this impacts a listener’s ability to understand the semantic and ture of a musical sequence involves playing the intended notes with
propositional content of the sentences. Recent research on this topic appropriate timing. Of course, this is not to say that a few wrong
has shown that spoken language comprehension is strikingly robust notes will ruin musical communication (even professionals make
to manipulations of pitch contour. Patel et al. (2010) measured the occasional mistakes in complex passages), but by and large the notes
intelligibility of natural Mandarin Chinese sentences with intact vs. and rhythms of a performance need to adhere fairly closely to a
flattened (monotone) pitch contours (the latter created via speech specified model (such as a musical score, or to a model provided
resynthesis, with pitch fixed at the mean fundamental frequency of by a teacher) for a performance to be deemed adequate. Even in
the sentence). Native speakers of Mandarin found the monotone improvisatory music such as jazz or the classical music of North
sentences just as intelligible as the natural sentences when heard India, a performer must learn to produce the sequence of notes
in a quiet background. Hence despite the complete removal of all they intend to play, and not other, unintended notes. Crucially, this
details of pitch variation, the sentences were fully intelligible. How requirement places fairly strict demands on the regulation of pitch
is this possible? Presumably listeners used the remaining phonetic and timing, because intended notes (whether from a musical score,
information and their knowledge of Mandarin to guide their per- a teacher’s model, or a sequence created “on the fly” in improvisa-
ception in a way that allowed them to infer which words were being tion) are never far in pitch or duration from unintended notes.
said. The larger point is that spoken language comprehension is For example, as mentioned above, a note just one semitone away
remarkably robust to lack of detail in pitch variation, and this pre- from an intended note can make a perceptually salient change in
sumably relaxes the demands placed on high-precision encoding a melody (e.g., when the note departs from the prevailing musical
of periodicity patterns. key). Similarly, a note that is just a few hundred milliseconds late
It seems likely that this sort of robustness is not just limited to compared to its intended onset time can make a salient change in
the acoustic feature of periodicity. One reason that spoken lan- the rhythmic feel of a passage (e.g., the difference between an “on
guage comprehension is so robust is that it involves integrating beat” note and a “syncopated” note). In short, conveying musi-
multiple cues, some of which provide redundant sources of infor- cal structure involves a high degree of precision, due to the fact
mation regarding a sound’s phonological category. For example, that listeners use fine acoustic details in judging the structure of
in judging whether a sound within a word is a /b/ or a /p/, multi- the music they are hearing. Furthermore, listeners use very fine
ple acoustic cues are relevant, including voice onset time (VOT), acoustic details in judging the expressive qualities of a performance.
vowel length, fundamental frequency (F0; i.e., “microintonational” Empirical research has shown that expressive (vs. deadpan) perfor-
perturbations of F0, which behave differently after voiced vs. voice- mances involve subtle, systematic modifications to the duration,
less stop consonants), and first and second formant patterns (i.e., intensity, and (in certain instruments) pitch of notes relative to

their nominal score-based values (Repp, 1992; Clynes, 1995; Palmer, This section has argued that music perception is often likely to
1997). Listeners are quite sensitive to these inflections and use them place higher demands on the encoding of certain acoustic features
in judging the emotional force of musical sequences (Bhatara et al., than does speech perception, at least in terms of what is needed for
2011). Indeed, neuroimaging research has shown that expressive adequate communication in the two domains. However, OPERA
performances containing such inflections are more likely (than makes no a priori assumption that influences between musical and
deadpan performances) to activate limbic and paralimbic brain linguistic neural encoding are unidirectional. Hence an important
areas associated with emotion processing (Chapin et al., 2010). question for future work is whether certain types of linguistic expe-
This helps explain why musical training involves learning to pay rience with heightened demands in terms of auditory processing
attention to the fine acoustic details of sound sequences and to (e.g., multilingualism, or learning a tone language) can impact the
control them with high-precision, since these details matter for the neural encoding of music (cf. Bidelman et al., 2011). This is an inter-
esthetic and emotional qualities of musical sequences. esting question, but is not explored in the current paper. Instead,
Returning to our example of pitch variation in speech vs. music, the focus is on explaining why musical training would benefit the
the above paragraph suggests that the adequate communication of neural encoding of speech, given the growing body of evidence
melodic music requires the detailed regulation and perception of for such benefits and the practical importance of such findings.
pitch patterns. According to OPERA, this puts higher demands on
the sensory encoding of periodicity than does speech, and helps Emotion, Repetition, and Attention
drive experience-dependent plasticity in subcortical networks For musical training to enhance the neural encoding of speech, the
that encode periodicity. Yet since speech and music share such musical activities that engage speech-processing networks must
networks, speech processing benefits (cf. the study of Wong et al., elicit strong positive emotion, be frequently repeated, and be asso-
2007, described in section The Auditory Brainstem Response to ciated with focused attention. These factors work in concert to
Speech: Origins and Plasticity). promote adaptive plasticity, and are discussed together here.
Yet would this benefit in the neural encoding of voice periodicity As noted in the previous section, music can place higher demands
have any consequences for real-world language skills? According to on the nervous system than speech does, in terms of the precision
the study of Patel et al. (2010) described above, the details of spo- of encoding of a particular acoustic feature. Yet this alone is not
ken pitch patterns are not essential for adequate spoken language enough to drive experience-dependent plasticity to enhance the
understanding, even in the tone language Mandarin. So why would encoding of that feature. There must be some (internal or external)
superior encoding of voice pitch patterns be helpful to a listener? motivation to enhance the encoding of that feature, and there must
One answer to this question is suggested by another experimental be sufficient opportunity to improve encoding over time. Musical
manipulation in the study of Patel et al. (2010). In this manipula- training has the potential to fulfill these conditions. Accurate music
tion, natural vs. monotone Mandarin sentences were embedded in performance relies on accurate perception of the details of sound,
background babble noise. In this case, the monotone sentences were which is presumably associated in turn with high-precision encod-
significantly less intelligible to native listeners (in the case where the ing of sound features. Within music, accurate performance is typi-
noise was as loud as the target sentence, monotone sentences were cally associated with positive emotion and reward, for example,
20% less intelligible than natural sentences). The precise reasons via internal satisfaction, praise from others, and from the pleasure
why natural pitch modulation enhances sentence intelligibility in of listening to well-performed music, in this case music produced
noise remain to be determined. Nevertheless, based on these results by oneself or the group one is in. (Note that according to this view,
it seems plausible that a nervous system which has high-precision the particular emotions expressed by the music one plays, e.g., joy,
encoding of vocal periodicity would have an advantage in under- sadness, tranquility, etc., are not crucial. Rather, the key issue is
standing speech in a noisy background, and recent empirical data whether the musical experience as a whole is emotionally reward-
support this conjecture (Song et al., 2010). Combining this finding ing.) Furthermore, accurate performance is typically acquired via
with the observation that musicians show superior brainstem encod- extensive practice, including frequent repetitions of particular
ing of voice F0 (Wong et al., 2007) leads to the idea that musically pieces. The neurobiological association between accurate perfor-
trained listeners should show better speech intelligibility in noise. mance and emotional rewards, and the opportunity to improve
In fact, it has recently been reported that musically trained with time (e.g., via extensive practice) create favorable conditions for
individuals show superior intelligibility for speech in noise, using promoting plasticity in the networks that encode acoustic features.
standard tests (Parbery-Clark et al., 2009). This study also examined Another factor likely to promote plasticity is focused attention
brainstem responses to speech, but rather than focusing on the on the details of sound during musical training. Animal studies of
encoding of periodicity in syllables with different pitch contour, it auditory training have shown that training-related cortical plastic-
examined the brainstem response to the syllable /da/ in terms of ity is facilitated when sounds are actively attended vs. experienced
latency, representation of speech harmonics, and overall response passively (e.g., Fritz et al., 2005; Polley et al., 2006). As noted by
morphology. When /da/ was heard in noise, all of these param- Jagadeesh (2006), attention “marks a particular set of inputs for
eters were enhanced in musically trained individuals compared to special treatment in the brain,” perhaps by activating cholinergic
their untrained counterparts. This suggests that musical training systems or increasing the synchrony of neural firing, and this modu-
influences the encoding of the temporal onsets and spectral details lation plays an important role in inducing neural plasticity (cf.
of speech, in addition to the encoding of voice periodicity, all of Kilgard and Merzenich, 1998; Thiel, 2007; Weinberger, 2007). While
which may contribute to enhanced speech perception in noise (cf. animal studies of attention and plasticity have focused on corti-
Chandrasekaran et al., 2009). cal circuits, it is worth recalling that the auditory system has rich

cortico-subcortical (corticofugal) connections, so that changes at

the cortical level have the potential to influence subcortical circuits,
and hence, the basic encoding of sound features (Schofield, 2010).
Focused attention on sound may also help resolve an apparent
“bootstrapping” problem raised by OPERA: if musical training is to
improve the precision of auditory encoding, how does this process
get started? For example, focusing on pitch, how does the auditory
system adjust itself to register finer pitch distinctions if it cannot
detect them in the first place? One answer might be that attention
activates more neurons in the frequency (and perhaps periodicity)
channels that encode pitch, such that more neurons are recruited to
deal with pitch, and hence acuity improves (P. Cariani, pers. comm.;
cf. Recanzone et al., 1993; Tervaniemi et al., 2009).
It is worth noting that the emotion, repetition, and attention
criteria of OPERA are falsifiable. Imagine a child who is given
weekly music lessons but who dislikes the music he or she is taught,
who does not play any music outside of lessons, and who is not
very attentive to the music during lessons. In such circumstances,
OPERA predicts that musical training will not result in enhanced
neural encoding of speech.
Putting OPERA to work: musical training and

linguistic reading skills
There is growing interest in links between musical training, audi-
tory processing, and linguistic reading skills. This stems from two
lines of research. The first line has shown a relationship between
musical abilities and reading skills in normal children, for exam- Figure 8 | Examples of non-linguistic tonal stimulus waveforms for rise
times of 15 (A) and 300 ms (B). From Goswami et al. (2002), reproduced
ple, via correlational and experimental studies (e.g., Anvari et al., with permission.
2002; Moreno et al., 2009). For example, Moreno et al. (2009)
assigned normal 8-year olds to 6 months of music vs. painting
lessons, and found that after musical (but not painting) training, in this range are necessary for speech intelligibility (Drullman et al.,
children showed enhanced reading abilities and improved auditory 1994; cf. Ghitza and Greenberg, 2009). In terms of connections to
discrimination in speech, with the latter shown by both behavioral reading, Goswami has suggested that problems in envelope per-
and neural measures (scalp-recorded cortical EEG). ception during language development could result in less robust
The second line of research has demonstrated that a substantial phonological representations at the syllable-level, which would
portion of children with reading problems have auditory processing then undermine the ability to consciously segment syllables into
deficits, leading researchers to wonder whether musical training individual speech sounds (phonemes). Phonological deficits are a
might be helpful for such children (e.g., Overy, 2003; Tallal and core feature of dyslexia (if the dyslexia is not primarily due to visual
Gaab, 2006; Goswami, 2010). For example, Goswami has drawn problems), likely because reading requires the ability to segment
attention to dyslexics’ impairments in discriminating the rate of words into individual speech sounds in order to map the sounds
change of amplitude envelope at a sound’s onset (its “rise time”; onto visual symbols. In support of Goswami’s ideas about relations
Figure 8), and have shown that such deficits predict reading abilities between envelope processing and reading, a recent meta-analysis of
even after controlling for the effects of age and IQ. Of course, rise- studies measuring dyslexics’ performance on non-speech auditory
time deficits are not the only auditory processing deficits that have tasks and on reading tasks reported that amplitude modulation and
been associated with dyslexia (Tallal and Gaab, 2006; Vandermosten rise time discrimination were linked to developmental dyslexia in
et al., 2010), but they are of interest to speech–music studies because 100% of the studies that they reviewed (Hämäläinen et al., in press).
they suggest a problem with encoding amplitude envelope patterns, Amplitude envelope is also an important acoustic feature of
and amplitude envelope is an important feature in both speech and musical sounds. Extensive pychophysical research has shown that
music perception. envelope is one of the major contributors to a sound’s musical tim-
In speech, the amplitude envelope is the relatively slow (syllable- bre (e.g., Caclin et al., 2005). For example, one of the important cues
level) modulation of overall energy within which rapid spectral that allows a listener to distinguish between the sounds of a flute
changes take place. Amplitude envelopes play an important role as and a French horn is the amplitude envelope of their musical notes
cues to speech rhythm (e.g., stress) and syllable boundaries, which (Strong and Clark, 1967). Furthermore, the slope of the amplitude
in turn help listeners segment words from the flow of speech (Cutler, envelope at tone onset is an important cue to the perceptual attack
1994). Envelope fluctuations in speech have most of their energy in time of musical notes, and thus to the perception of rhythm and
the 3–20 Hz range, with a peak around 5 Hz (Greenberg, 2006), and timing in sequences of musical events (Gordon, 1987). Given that
experimental manipulations have shown that envelope modulations envelope is an important acoustic feature in both speech and music,

the OPERA hypothesis leads to the prediction that musical training light of Goswami’s ideas, described above, it is worth noting that
which relies on high-precision envelope processing could benefit in a subsequent study Abrams et al. [2009] repeated their study
the neural processing of speech envelopes, via mechanisms of adap- with good vs. poor readers, aged 9–15 years. The good readers
tive neural plasticity, if the five conditions of OPERA are met. Hence showed a strong right-hemisphere advantage for envelope tracking
the rest of this section is devoted to examining these conditions in in multiple measures [e.g., higher cross-correlation, shorter-latency
terms of envelope processing in speech and music. between neural response and speech input], while poor readers did
The first condition is that envelope processing in speech and not. Furthermore, empirical measures of neural envelope tracking
music relies on overlapping neural circuitry. What is known about predicted reading scores, even after controlling for IQ differences
amplitude envelope processing in the brain? Research on this between good and poor readers).
question in humans and other primates has focused on cortical Thus it appears that right-hemisphere auditory cortical regions
responses to sound. It is very likely, however, that subcortical circuits (and very likely, subcortical inputs to these regions) are involved
are involved in envelope encoding, so that measures of envelope in envelope processing in speech. Are the same circuits involved in
processing taken at the cortex reflect in part the encoding capacities envelope processing in music? This remains to be tested directly,
of subcortical circuits. Yet since extant primate data come largely and could be studied using an individual differences approach.
from cortical studies, those will be the focus here. For example, one could use the design of Abrams et al. (2008)
Primate neurophysiology research suggests that the temporal to examine cortical responses to the amplitude envelope of spo-
envelope of complex sound is represented by population-level ken sentences and musical melodies in the same listeners, with
activity of neurons in auditory cortex (Nagarajan et al., 2002). In the melodies played using instruments that have sustained (vs.
human auditory research, two theories suggest that slower temporal percussive) notes, and hence envelopes reminiscent of speech syl-
modulations in sounds (such as amplitude envelope) are preferen- lables. If envelope processing mechanisms are shared between
tially processed by right-hemisphere cortex (Zatorre et al., 2002; speech and music, then one would predict that individuals would
Poeppel, 2003). Prompted by these theories, Abrams et al. (2008) show similar envelope-tracking quality across the two domains.
recently examined cortical encoding of envelope using scalp EEG. While such data are lacking, there are indirect indications of a
They collected neural responses to an English sentence (“the young relationship between envelope processing in music and speech.
boy left home”) presented to healthy 9- to 13-year-old children. In music, one set of abilities that should depend on envelope
(The children heard hundreds of repetitions of the sentence in one processing are rhythmic abilities, because such abilities depend
ear, while watching a movie and hearing the soundtrack quietly on sensitivity to the timing of musical notes, and envelope is an
in the other ear.) The EEG response to the sentence was averaged important cue for the perceptual onset and duration of event
across trials and low-pass filtered at 40 Hz to focus on cortical onsets in music (Gordon, 1987). Hence if musical and linguistic
responses and how they might reflect the amplitude envelope of the amplitude envelopes are processed by overlapping brain circuits,
sentence. Abrams et al. discovered that the EEG signal at temporal one would expect (somewhat counterintuitively) a relationship
electrodes over both hemispheres tracked the amplitude envelope between musical rhythmic abilities and reading abilities. Is there
of the spoken sentence. (The authors argue that the neural EEG any evidence for such a relationship? In fact, Goswami and col-
signals they studied are likely to arise from activity in secondary leagues have reported that dyslexics have problems with musical
auditory cortex.) Notably, the quality of tracking, as measured by rhythmic tasks (e.g., Thompson and Goswami, 2008; Huss et al.,
cross-correlating the EEG waveform with the speech envelope, was 2011), and have shown that their performance on these tasks
far superior in right-hemisphere electrodes (approx. 100% supe- correlates with their reading skills, after controlling for age and
rior), in contrast to the usual left hemisphere dominance for spoken IQ. Furthermore, normal 8-year-old children show positive cor-
language processing (Hickok and Poeppel, 2007; Figure 9). (In relations between performance on rhythm discrimination tasks
(but not pitch discrimination tasks) and reading tasks, even after
factoring out effects of age, parental education, and the num-
ber of hours children spend reading per week (Corrigall and
Trainor, 2010).
Based on such findings, let us assume for the sake of argument
that the first condition of OPERA is satisfied, that is, that envelope
processing in speech and music utilizes overlapping brain networks.
It is not required that such networks are entirely overlapping, simply
that they overlap to a significant degree at some level of processing
(e.g., subcortical, cortical, or both). With such an assumption, one
can then turn to the next component of OPERA and consider what
sorts of musical tasks would promote high-precision envelope pro-
Figure 9 | Grand average cortical responses from temporal electrodes cessing. In ordinary musical circumstances, the amplitude envelope
T3 and T4 (red: right hemisphere, blue: left hemisphere) and broadband of sounds is an acoustic feature relevant to timbre, and while timbre
speech envelope (black) for the sentence “the young boy left home.” is an important attribute of musical sound, it (unlike pitch) is rarely
Ninety-five milliseconds of the prestimulus period is plotted. The speech a primary structural parameter for musical sequences, at least in
envelope was shifted forward in time 85 ms to enable comparison to cortical
Western melodic music (Patel, 2008). Hence in order to encourage
responses. From Abrams et al. (2008), reproduced with permission.
high-precision envelope processing in music, it may be necessary

to devise novel musical activities, rather than relying on standard is important to note that the OPERA hypothesis is not proscriptive:
musical training. For example, using modern digital technology in it says nothing about the relative merits of musical vs. linguistic
which keyboards can produce any type of synthesized sounds, one training for speech sound encoding. Instead, it tries to account for
could create synthesized sounds with similar spectral content but why musical training would benefit speech sound encoding in the
slightly different amplitude envelope patterns, and create musical first place, given the growing empirical evidence that this is indeed
activities (e.g., composing, playing, listening) which rely on the the case. Hence the relative efficacy of musical vs. linguistic train-
ability to distinguish such sounds. Note that such sounds need ing for speech sound encoding is an empirical question which can
not be musically unnatural: acoustic research on orchestral wind- only be resolved by direct comparison in future studies. A strong
instrument tones has shown, for example, that the spectrum of motivation for conducting such comparisons is the demonstra-
the flute, bassoon, trombone, and French horn are similar, and tion that non-linguistic auditory perceptual training generalizes
that listeners rely on envelope cues in distinguishing between these to linguistic discrimination tasks that rely on acoustic cues similar
instruments (Strong and Clark, 1967). The critical point is that suc- to those trained in the non-linguistic context (Lakshimnarayanan
cess at the musical activity should require high-precision processing and Tallal, 2007).
of envelope patterns. While direct comparisons of musical vs. linguistic training
For example, if working with young children, one could create have yet to be conducted, it is worth considering some of the
computer-animated visual characters who “sing” with different potential merits of music-based training. First, musical activi-
“voices” (i.e., their “songs” are non-verbal melodies in which all ties are often very enjoyable, reflecting the rich connections
tones have a particular envelope shape). After learning to associ- between music processing and the emotion systems of the brain
ate different visual characters and their voices, one could then do (Koelsch, 2010; Salimpoor et al., 2011). Hence it may be easier
games where novel melodies are heard without visual cues and to have individuals (especially children) participate repeatedly
the child has to guess which character is singing. Such “envelope in training tasks, particularly in home-based tasks which require
training” games could be done adaptively, so that at the start of voluntary participation. Second, if an individual is experiencing
training, the envelope shapes of the different characters are quite a language problem, then a musical activity may not carry any
different, and then once successful discrimination is achieved, new of the negative associations that have developed around the lan-
characters are introduced whose voices are more similar in terms guage deficits and language-based tasks. This increases the chances
of envelope cues. of associating auditory training with strong positive emotions,
According to OPERA, if this sort of training is to have any effect which in turn may facilitate neural plasticity (as noted in section
on the precision of envelope encoding by the brain, the musical Emotion, Repetition, and Attention). Third, speech is acoustically
tasks should be associated with strong positive emotion, extensive complex, with many acoustic features varying at the same time
repetition, and focused attention. Fortunately, music processing is (e.g., amplitude envelope, harmonic spectrum), and its perception
known to have a strong relationship to the brain’s emotion systems automatically engages semantic processes that attempt to extract
(Koelsch, 2010) and it seems plausible that the musical tasks could conceptual meaning from the signal, which draws attention away
be made pleasurable (e.g., via the use of attractive musical sounds from the acoustic details of the signal. Musical sounds, in contrast,
and melodies, and stimulating rewards for good performance). can be made acoustically relatively simple, with variation primarily
Furthermore, if they are sufficiently challenging, then participants occurring along specific dimensions that are the focus of training.
will likely want to engage in them repeatedly and with focused For example, if the focus is on training sensitivity to amplitude
attention. envelope (cf. section Putting OPERA to work: musical training
To summarize, the OPERA hypothesis predicts that musical and linguistic reading skills), sounds can be created in which the
training which requires high-precision amplitude envelope pro- spectral content is simple and stable, and in which the primary
cessing will benefit the neural encoding of amplitude envelopes in differences between tones are in envelope structure. Furthermore,
speech, via mechanisms of neural plasticity, if the five conditions of musical sounds do not engage semantic processing, leaving the
OPERA are met. Based on research showing relationships between perceptual system free to focus more attention on the details of
envelope processing and reading abilities (e.g., Goswami, 2010), sound. This ability of musical sounds to isolate particular features
this in turn may benefit linguistic reading skills. for attentive processing could lower the overall complexity of audi-
tory training tasks, and hence make it easier for individual to make
Musical vs. linguistic training for speech sound rapid progress in increasing their sensitivity to such features, via
encoding mechanisms of neural plasticity.
As described in the previous section, the OPERA hypothesis leads A final merit of musical training is that it typically involves build-
to predictions for how musical training might benefit specific ing strong sensorimotor links between auditory and motor skills
linguistic abilities via enhancements of the neural encoding of (i.e., the sounds one produces are listened to attentively, in order
speech. Yet this leads to an obvious and important question. Why to adjust performance to meet a desired model). Neuroscientific
attempt to improve the neural encoding of speech by training in research suggests that sensorimotor musical training is a stronger
other domains? If one wants to improve speech processing, would driver of neural plasticity in auditory cortex than purely auditory
it not be more effective to do acoustic training in the context musical training (Lappe et al., 2008). Hence musical training pro-
of speech? Indeed, speech-based training programs that aim to vides an easy, ecologically natural route to harness the power of
improve sensitivity to acoustic features of speech are now widely sensorimotor processing to drive adaptive neural plasticity in the
used (e.g., the Fast ForWord program, cf. Tallal and Gaab, 2006). It auditory system.

Conclusion digital instruments may be necessary, which allow controlled

The OPERA hypothesis suggests that five essential conditions manipulation of sound features. Thus OPERA raises the idea
must be met in order for musical training to drive adaptive plas- that in the future, musical activities aimed at benefiting speech
ticity in speech processing networks. This hypothesis generates processing should be purposely shaped to optimize the effects
specific predictions, for example, regarding the kinds of musi- of musical training.
cal training that could benefit reading skills. It also carries an From a broader perspective, OPERA contributes to the growing
implication regarding the notion of “musical training.” Musical body of research aimed at understanding the relationship between
training can involve different skills depending on what instru- musical and linguistic processing in the human brain (Patel, 2008).
ment and what aural abilities are being trained. (For example, Understanding this relationship will likely have significant implica-
learning to play the drums places very different demands on the tions for how we study and treat a variety of language disorders,
nervous system than learning to play the violin.) The OPERA ranging from sensory to syntactic processing (Jentschke et al., 2008;
hypothesis states that the benefits of musical training depend Patel et al., 2008).
on the particular acoustic features emphasized in training, the
demands that music places on those features in terms of the preci- Acknowledgments
sion of processing, and the degree of emotional reward, repetition Supported by Neurosciences Research Foundation as part of its
and attention associated with musical activities. According to this research program on music and the brain at The Neurosciences
hypothesis, simply giving an individual music lessons may not Institute, where ADP is the Esther J. Burnham Senior Fellow. I
result in any benefits for speech processing. Indeed, depending thank Dan Abrams, Peter Cariani, Bharath Chandrasekaran, Jeff
on the acoustic feature being trained (e.g., amplitude envelope), Elman, Lori Holt, John Iversen, Nina Kraus, Stephen McAdams,
learning a standard musical instrument may not be an effective Stefanie Shattuck-Hufnagel, and the reviewers of this paper for
way to enhance neural processing of that feature. Instead, novel insightful comments.
References Bidelman, G. M., Gandour, J. T., and in children,” in Paper presented at the developmental dyslexia: a new
Abrams, D. A., Nicol, T., Zecker, S., and Krishnan, A. (2011). Cross-domain 11th Intl. Conf. on Music Perception & hypothesis. Proc. Natl. Acad. Sci.
Kraus, N. (2008). Right-hemisphere effects of music and language expe- Cognition (ICMPC11), August 2010, U.S.A. 99, 10911–10916.
auditory cortex is dominant for coding rience on the representation of pitch Seattle, WA. Goswami, U. (2010). A temporal sam-
syllable patterns in speech. J. Neurosci. in the human auditory brainstem. J. Cutler, A. (1994). Segmentation problems, pling framework for developmental
28, 3958–3965. Cogn. Neurosci. 23, 425–434. rhythmic solutions. Lingua 92, 81–104. dyslexia. Trends Cogn. Sci. 15, 3–10.
Abrams, D. A., Nicol, T., Zecker, S., and Caclin, A., McAdams, S., Smith, B. K., and Drullman, R., Festen, J. M., and Plomp, Greenberg, S. (2006). “A multi-tier frame-
Kraus, N. (2009). Abnormal corti- Winsberg, S. (2005). Acoustic corre- R. (1994). Effect of temporal enve- work for understanding spoken lan-
cal processing of the syllable rate of lates of timbre space dimensions: a lope smearing on speech reception. J. guage,” in Listening to Speech: An
speech in poor readers. J. Neurosci. 29, confirmatory study using synthetic Acoust. Soc. Am. 95, 1053–1064. Auditory Perspective, eds S. Greenberg
7686–7693. tones. J. Acoust. Soc. Am. 118, 471–482. Edelman, G. M., and Gally, J. (2001). and W. A. Ainsworth (Mahwah, NJ:
Ahissar, M., Nahum, M., Nelken, I., and Cariani, P. A., and Delgutte, B. (1996). Degeneracy and complexity in bio- Erlbaum), 411–433.
Hochstein, S. (2009). Reverse hier- Neural correlates of the pitch of com- logical systems. Proc. Natl. Acad. Sci. Hämäläinen, J. A., Salminen, H. K., and
archies and sensory learning. Philos. plex tones. I. Pitch and pitch salience. J. U.S.A. 98, 13763–13768. Leppänen, P. H. T. (in press). Basic
Trans. R. Soc. Lond. B Biol. Sci. 364, Neurophysiol. 76, 1698–1716. Eerola, T., Himberg, T., Toiviainen, P., auditory processing deficits in dys-
285–299. Chandrasekaran, B., Hornickel, J., Skoe, and Louhivuori, J. (2006). Perceived lexia: Review of the behavioural,
Anvari, S., Trainor, L. J., Woodside, J., and E., Nicol, T., and Kraus, N. (2009). complexity of western and African event-related potential and magne-
Levy, B. A. (2002). Relations among Context-dependent encoding in the folk melodies by western and African toencephalographic evidence. J. Learn.
musical skills, phonological process- human auditory brainstem relates to listeners. Psychol. Music 34, 337–371. Disabil.
ing, and early reading ability in pre- hearing speech in noise: implications Ferreira, F., and Patson, N. D. (2007). Hickok, G., and Poeppel, D. (2007). The
school children. J. Exp. Child Psychol. for developmental dyslexia. Neuron The ‘good enough’ approach to lan- cortical organization of speech pro-
83, 111–130. 64, 311–319. guage comprehension. Lang. Linguist. cessing. Nat. Rev. Neurosci. 8, 393–402.
Bajo, V. M., Nodal, F. R., Moore, D. Chandrasekaran, B., and Kraus, N. Compass 1, 71–83. Holt, L. L., and Idemaru, K. (2011).
R., and King, A. J. (2010). The (2010). The scalp-recorded brain- Fritz, J., Elhilali, M., and Shamma, S. “Generalization of dimension-based
descending corticocollicular path- stem response to speech: neural ori- (2005). Active listening: task-depend- statistical learning of speech,” in Proc.
way mediates learning-induced gins and plasticity. Psychophysiology ent plasticity of spectrotemporal 17th Intl. Cong. Phonetic Sci. (ICPhS
auditory plasticity. Nat. Neurosci. 47, 236–246. receptive fields in primary auditory XVII), Hong Kong, China.
13, 252–260. Chapin, H., Jantzen, K., Kelso, S. J. A., cortex. Hear. Res. 206, 159–176. Huss, M., Verney, J. P., Fosker, T., Mead,
Banai, K., Hornikel, J., Skoe, E., Nicol, Steinberg, F., and Large, E. (2010). Ghitza, O., and Greenberg, S. (2009). On N., and Goswami, U. (2011). Music,
T., Zecker, S., and Kraus, N. (2009). Dynamic emotional and neural the possible role of brain rhythms in rhythm, rise time perception and
Reading and subcortical auditory responses to music depend on per- speech perception: Intelligibility of developmental dyslexia: perception
function. Cereb. Cortex 19, 2699–2707. formance expression and listener time compressed speech with periodic of musical meter predicts reading
Bendor, D., and Wang, X. (2005). The experience. PLoS ONE 5: e13812. doi: and aperiodic insertions of silence. and phonology. Cortex 47, 674–689.
neuronal representation of pitch in 10.1371/journal.pone.0013812 Phonetica 66, 113–126. Hyde, K. L., Lerch, J., Norton, A., Forgeard,
primate auditory cortex. Nature 436, Clynes, M. (1995). Microstructural musi- Gordon, J. W. (1987). The perceptual M., Winner, E., Evans, A. E., and
1161–1165. cal linguistics: composers’ pulses are attack time of musical tones. J. Acoust. Schlaug, G. (2009). Musical training
Bhatara, A., Tirovolas, A. K., Duan, L. liked most by the best musicians. Soc. Am. 82, 88–105. shapes structural brain development.
M., Levy, B., and Levitin, D. J. (2011). Cognition 55, 269–310. Goswami, U., Thompson, J., Richardson, J. Neurosci. 29, 3019–3025.
Perception of emotional expression in Corrigall, K., and Trainor, L. (2010). “The U., Stainthorp, R., Hughes, D., Jagadeesh, B. (2006). “Attentional modu-
musical performance. J. Exp. Psychol. predictive relationship between length Rosen, S., and Scott, S. K. (2002). lation of cortical plasticity,” in Textbook
Hum. Percept. Perform. 37, 921–934. of musical training and cognitive skills Amplitude envelope onsets and of Neural Repair and Rehabilitation:

Neural Repair and Plasticity, Vol. Exp. Psychol. Hum. Percept. Perform. Patel, A. D., Xu, Y., and Wang, B. (2010). Strait, D. L., Kraus, N., Parbery-Clark, A.,
1. eds M. Selzer, S. E. Clarke, L. G. 34, 1609–1631. “The role of F0 variation in the intel- and Ashley, R. (2010). Musical expe-
Cohen, P. W. Duncan, and F. H. Gage Moreno, S., Marques, C., Santos, ligibility of Mandarin sentences,” in rience shapes top-down auditory
(Cambridge: Cambridge University A., Santos, M., Castro, S. L., and Proc. Speech Prosody 2010, May 11–14, mechanisms: evidence from masking
Press), 194–205. Besson, M. (2009). Musical train- 2010, Chicago, IL, USA. and auditory attention performance.
Jentschke, S., Koeslch, S., Sallat, S., and ing influences linguistic abilities in Peretz, I., and Coltheart, M. (2003). Hear. Res. 261, 22–29.
Friederici, A. (2008). Children with 8-year-old children: more evidence Modularity of music processing. Nat. Strait, D. L., Skoe, E., Kraus, N., and
specific language impairment also for brain plasticity. Cereb. Cortex 19, Neurosci. 6, 688–691. Ashley, R. (2009). Musical experi-
show impairment of music-syntactic 712–723. Peretz, I., Cummings, S., and Dubé, M-P. ence and neural efficiency: effects of
processing. J. Cogn. Neurosci. 20, Musacchia, G., Sams, M., Skoe, E., and (2007). The genetics of congenital training on subcortical processing of
1940–1951. Kraus, N. (2007). Musicians have amusia (or tone-deafness): a family vocal expressions of emotion. Eur. J.
Kaas, J., and Hackett, T. (2000). enhanced subcortical auditory and aggregation study. Am. J. Hum. Genet. Neurosci. 29, 661–668.
Subdivisions of auditory cortex audiovisual processing of speech and 81, 582–588. Strong, W., and Clark, M. C. (1967).
and processing streams in primates. music. Proc. Natl. Acad. Sci. U.S.A. 104, Poeppel, D. (2003). The analysis of speech Perturbations of synthetic orchestral
Proc. Natl. Acad. Sci. U.S.A. 97, 15894–15898. in different temporal integration wind-instrument tones. J. Acoust. Soc.
11793–11799. Musacchia, G., Strait, D. L., and Kraus, N. windows: cerebral lateralization as Am. 41, 277–285.
Koelsch, S. (2010). Towards a neural basis (2008). Relationships between behav- ‘asymmetric sampling in time’. Speech Suga, N. (2008). Role of corticofugal
of music-evoked emotions. Trends ior, brainstem and cortical encoding of Commun. 41, 245–255. feedback in hearing. J. Comp. Physiol.
Cogn. Sci. 14, 131–137. seen and heard speech in musicians Polley, D. B., Steinberg, E. E., and A Neuroethol. Sens. Neural Behav.
Kilgard, M., and Merzenich, M. (1998). and nonmusicians. Hear. Res. 241, Merzenich, M. M. (2006). Perceptual Physiol. 194, 169–183.
Cortical reorganization enabled by 34–42. learning directs auditory cortical Tallal, P., and Gaab, N. (2006). Dynamic
nucleus basalis activity. Science 279, Nagarajan, S. S., Cheung, S. W., map reorganization through top- auditory processing, musical expe-
1714–1718. Bedenbaugh, P., Beitel, R. E., down influences. J. Neurosci. 26, rience and language development.
Kral, A., and Eggermont, J. (2007). What’s Schreiner, C. E., and Merzenich, M. M. 4970–4982. Trends Neurosci. 29, 382–390.
to lose and what’s to learn: develop- (2002). Representation of spectral and Recanzone, G. H., Schreiner, C. C., and Tervaniemi, M., Kruck, S., De Baene, W.,
ment under auditory deprivation, temporal envelope of twitter vocaliza- Merzenich, M. M. (1993). Plasticity Schröger, E., Alter, K., and Friederici,
cochlear implants and limits of cor- tions in common marmoset primary in the frequency representation of A. D. (2009). Top-down modulation
tical plasticity. Brain Res. Rev. 56, auditory cortex. J. Neurophysiol. 87, primary auditory cortex following of auditory processing: effects of
259–269. 1723–1737. discrimination training in adult owl sound context, musical expertise and
Kraus, N., and Chandrasekaran, B. (2010). Overy, K. (2003). Dyslexia and music: monkeys. J. Neurosci. 13, 87–103. attentional focus. Eur. J. Neurosci. 30,
Music training for the development from timing deficits to musical inter- Repp, B. H. (1992). Diversity and com- 1636–1642.
of auditory skills. Nat. Rev. Neurosci. vention. Ann. N. Y. Acad. Sci. 999, monality in music performance: an Thiel, C. M. (2007). Pharmacological
11, 599–605. 497–505. analysis of timing microstructure in modulation of learning-induced
Krishnan, A., Xu, Y., Gandour, J., and Palmer, C. (1997). Music performance. Schumann’s “Träumerei.” J. Acoust. plasticity in human auditory cortex.
Cariani, P. (2005). Encoding of pitch Ann. Rev. Psychol. 48, 115–138. Soc. Am. 92, 2546–2568. Restor. Neurol. Neurosci. 25, 435–443.
in the human brainstem is sensitive to Panizzon, M. W., Fennema-Notestine, Russo, N., Nicol, T., Musacchia, G., and Thompson, J. M., and Goswami, U.
language experience. Brain Res. Cogn. C., Eyler, L. T., Jernigan, T. L., Prom- Kraus, N. (2004). Brainstem response (2008). Rhythmic processing in chil-
Brain Res. 25, 161–168. Wormley, E., Neale, M., Jacobson, K., to speech syllables. Clin. Neurophysiol. dren with developmental dyslexia:
Lakshimnarayanan, K., and Tallal, P. Lyons, M. J., Grant, M. D., Franz, C. 115, 2021–2030. auditory and motor rhythms link to
(2007). Generalization of non-lin- E., Xian, H., Tsuang, M., Tsuang, M., Salimpoor, V., Benovoy, M., Larcher, K., reading and spelling. J. Physiol. Paris
guistic auditory perceptual training to Fischl, B., Seidman, L., Dale, A., and Dagher, A., and Zatorre, R. (2011). 102, 120–129.
syllable discrimination. Restor. Neurol. Kremen, W. S. (2009). Distinct genetic Anatomically distinct dopamine Toscano, J. C., and McMurray, B. (2010).
Neurosci. 25, 263–272. influences on cortical surface area and release during anticipation and expe- Cue integration with categories:
Lappe, C., Herholz, S. C., Trainor, L. J., cortical thickness. Cereb. Cortex 19, rience of peak emotion to music. Nat. weighting acoustic cues in speech
and Pantev, C. (2008). Cortical plastic- 2728–2735. Neurosci. 14, 257–262. using unsupervised learning and
ity induced by short-term unimodal Parbery-Clark, A., Skoe, E., and Kraus, Schnupp, J., Nelken, I., and King, A. distributional statistics. Cogn. Sci. 34,
and multimodal musical training. J. N. (2009). Musical experience limits (2011). Auditory Neuroscience: Making 434–464.
Neurosci. 8, 9632–9639. the degradative effects of background Sense of Sound. Cambridge, MA: MIT Toscano, J. C., McMurray, B., Dennhardt,
Lee, K. M., Skoe, E., Kraus, N., and noise on the neural processing of Press. J., and Luck, S. J. (2010). Continuous
Ashley, R. (2009). Selective subcor- sound. J. Neurosci. 29, 14100–14107. Schofield, B. R. (2010). Projections from perception and graded categoriza-
tical enhancement of musical inter- Parbery-Clark, A., Strait, D. L., Anderson, auditory cortex to midbrain choliner- tion: electrophysiological evidence
vals in musicians. J. Neurosci. 29, S., Hittner, E., and Kraus, N. (2011). gic neurons that project to the inferior for a linear relationship between
5832–5840. Musical training and the aging audi- colliculus. Neuroscience 166, 231–240. the acoustic signal and perceptual
Liu, F., Patel, A. D., Fourcin, A., and tory system: implications for cognitive Skoe, E., and Kraus, N. (2010). Auditory encoding of speech. Psychol. Sci. 21,
Stewart, L. (2010). Intonation process- abilities and hearing speech in noise. brainstem response to complex 1532–1540.
ing in congenital amusia: discrimina- PLoS ONE 6: e18082. doi: 10.1371/ sounds: a tutorial. Ear Hear. 31, Tzounopoulos, T., and Kraus, N. (2009).
tion, identification, and imitation. journal.pone.0018082 302–324. Learning to encode timing: mecha-
Brain 133, 1682–1693. Patel, A. D. (2008). Music, Language, Song, J., Skoe, E., Banai, K., and Kraus, N. nisms of plasticity in the auditory
Mattys, S. L., White, L., and Melhorn, J. F. and the Brain. New York: Oxford (2010). Perception of speech in noise: brainstem. Neuron 62, 463–469.
(2005). Integration of multiple speech University Press. neural correlates. J. Cogn. Neurosci. 23, Vandermosten, M., Boets, B., Luts, H.,
segmentation cues: a hierarchical Patel, A. D., and Iversen, J. R. (2007). The 2268–2279. Poelmans, H., Golestani, N., Wouters,
framework. J. Exp. Psychol. General linguistic benefits of musical abilities. Song, J. H., Skoe, E., Wong, P. C., and J., and Ghesquière, P. (2010). Adults
134, 477–500. Trends Cogn. Sci. 11, 369–372. Kraus, N. (2008). Plasticity in the adult with dyslexia are impaired in categoriz-
McMurray, B., Aslin, R., Tanenhaus, M., Patel, A. D., Iversen, J. R., Wassenaar, human auditory brainstem following ing speech and nonspeech sounds on
Spivey, M., and Subik, D. (2008). M., and Hagoort, P. (2008). Musical short-term linguistic training. J. Cogn. the basis of temporal cues. Proc. Natl.
Gradient sensitivity to within- syntactic processing in agrammatic Neurosci. 10, 1892–1902. Acad. Sci. U.S.A. 107, 10389–10394.
category variation in speech: impli- Broca’s aphasia. Aphasiology 22, Stevens, K. N. (1998). Acoustic Phonetics. Vos, P. G., and Troost, J. M. (1989).
cations for categorical perception. J. 776–789. Cambridge, MA: MIT Press. Ascending and descending melodic

intervals: statistical findings and their experience shapes human brainstem Conflict of Interest Statement: The Front. Psychology 2:142. doi: 10.3389/
perceptual relevance. Music Percept. encoding of linguistic pitch patterns. author declares that the research was con- fpsyg.2011.00142
6, 383–396. Nat. Neurosci. 10, 420–422. ducted in the absence of any commercial This article was submitted to Frontiers in
Weinberger, N. (2007). Auditory asso- Zatorre, R. J., Belin, P., and Penhune, V. or financial relationships that could be Auditory Cognitive Neuroscience, a spe-
ciative memory and representational B. (2002). Structure and function of construed as a potential conflict of interest. cialty of Frontiers in Psychology.
plasticity in the primary auditory cor- auditory cortex: music and speech. Copyright © 2011 Patel. This is an open-
tex. Hear. Res. 229, 54–68. Trends Cogn. Sci. 6, 37–46. Received: 16 March 2011; paper pending access article subject to a non-exclusive license
Winer, J. (2006). Decoding the auditory Zatorre, R. J., and Gandour, J. T. (2008). published: 05 April 2011; accepted: 12 June between the authors and Frontiers Media
corticofugal systems. Hear. Res. 212, Neural specializations for speech and 2011; published online: 29 June 2011. SA, which permits use, distribution and
1–8. pitch: moving beyond the dichoto- Citation: Patel AD (2011) Why would reproduction in other forums, provided the
Wong, P. C., Skoe, E., Russo, N. M., Dees, mies. Phil. Trans. R. Soc. B 363, musical training benefit the neural encod- original authors and source are credited and
T., and Kraus, N. (2007). Musical 1087–1104. ing of speech? The OPERA hypothesis. other Frontiers conditions are complied with.

Why Would Musical Training Benefit The Neural Encoding of Speech? The OPERA Hypothesis

Uploaded by

Copyright:

Available Formats

Why Would Musical Training Benefit The Neural Encoding of Speech? The OPERA Hypothesis

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Why Would Musical Training Benefit The Neural Encoding of Speech? The OPERA Hypothesis

Uploaded by

Copyright:

Available Formats

Hypothesis and Theory Article

published: 29 June 2011

Why would musical training benefit the neural encoding of

Introduction can be enhanced by non-linguistic auditory training (i.e., learning

www.frontiersin.org June 2011 | Volume 2 | Article 142 | 1

Prior to these core sections of the paper, section “The Auditory

The auditory brainstem response to speech: origins

Frontiers in Psychology | Auditory Cognitive Neuroscience June 2011 | Volume 2 | Article 142 | 2

Figure 2 | Schematic diagram of the auditory system, illustrating the

As noted previously, the quality of this encoding is correlated with

www.frontiersin.org June 2011 | Volume 2 | Article 142 | 3

Frontiers in Psychology | Auditory Cognitive Neuroscience June 2011 | Volume 2 | Article 142 | 4

www.frontiersin.org June 2011 | Volume 2 | Article 142 | 5

Frontiers in Psychology | Auditory Cognitive Neuroscience June 2011 | Volume 2 | Article 142 | 6

www.frontiersin.org June 2011 | Volume 2 | Article 142 | 7

Frontiers in Psychology | Auditory Cognitive Neuroscience June 2011 | Volume 2 | Article 142 | 8

cortico-subcortical (corticofugal) connections, so that changes at

Putting OPERA to work: musical training and

www.frontiersin.org June 2011 | Volume 2 | Article 142 | 9

Frontiers in Psychology | Auditory Cognitive Neuroscience June 2011 | Volume 2 | Article 142 | 10

www.frontiersin.org June 2011 | Volume 2 | Article 142 | 11

Conclusion digital instruments may be necessary, which allow controlled

Frontiers in Psychology | Auditory Cognitive Neuroscience June 2011 | Volume 2 | Article 142 | 12

www.frontiersin.org June 2011 | Volume 2 | Article 142 | 13

Frontiers in Psychology | Auditory Cognitive Neuroscience June 2011 | Volume 2 | Article 142 | 14

You might also like