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J Exp Psychol Gen. Author manuscript; available in PMC 2019 April 24.
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Published in final edited form as:

J Exp Psychol Gen. 2017 April ; 146(4): 495–511. doi:10.1037/xge0000281.

Revisiting the ‘enigma’ of musicians with dyslexia: auditory

sequencing and speech abilities
Jennifer Zuk1,2,*, Paula Bishop-Liebler3,*, Ola Ozernov-Palchik1,4, Emma Moore5, Katie
Overy5, Graham Welch3, and Nadine Gaab1,2,6,+
1BostonChildren’s Hospital, Developmental Medicine Center, Laboratories of Cognitive
Neuroscience, Boston, MA, USA
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2Harvard Medical School, Boston, MA, USA

3International
Music Education Research Centre, Institute of Education, University College
London, London, UK
4Eliot-Pearson Department of Child Development, Tufts University, Somerville, MA, USA
5Institute of Music in Human and Social Development, Reid School of Music, Edinburgh, UK
6Harvard Graduate School of Education, Cambridge, MA, USA

Abstract
Previous research has suggested a link between musical training and auditory processing skills.
Musicians have shown enhanced perception of auditory features critical to both music and speech,
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suggesting that this link extends beyond basic auditory processing. It remains unclear to what
extent musicians who also have dyslexia show these specialized abilities, considering often-
observed persistent deficits that coincide with reading impairments. The present study evaluated
auditory sequencing and speech discrimination in 52 adults comprised of musicians with dyslexia,
nonmusicians with dyslexia, and typical musicians. An auditory sequencing task measuring
perceptual acuity for tone sequences of increasing length was administered. Furthermore, subjects
were asked to discriminate synthesized syllable continua varying in acoustic components of
speech necessary for intra-phonemic discrimination, which included spectral (formant frequency)
and temporal (voice onset time (VOT) and amplitude envelope) features. Results indicate that
musicians with dyslexia did not significantly differ from typical musicians and performed better
than nonmusicians with dyslexia for auditory sequencing as well as discrimination of spectral and
VOT cues within syllable continua. However, typical musicians demonstrated superior
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performance relative to both groups with dyslexia for discrimination of syllables varying in
amplitude information. These findings suggest a distinct profile of speech processing abilities in
musicians with dyslexia, with specific weaknesses in discerning amplitude cues within speech.
Since these difficulties seem to remain persistent in adults with dyslexia despite musical training,

This manuscript is not the copy of record and may not exactly replicate the final, authoritative version of the article. Please do not
copy or cite without authors’ permission.
+
Correspondence to: Nadine Gaab, Ph.D., Laboratories of Cognitive Neuroscience, Boston Children’s Hospital, Harvard Medical
School, 1 Autumn Street, Boston, MA 02215, Phone: (857) 218-3021, Fax: (617) 730-0518, [email protected].
*These authors contributed equally to this work.
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.
 Zuk et al. Page 2

this study only partly supports the potential for musical training to enhance the auditory
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processing skills known to be crucial for literacy in individuals with dyslexia.

Keywords
dyslexia; music; auditory; speech; children

Learning to read is essential for academic and vocational success, yet approximately 5–17%
of the population significantly struggles to read and comprehend text because of a specific
learning disorder known as dyslexia (Lyon, 2003; Peterson & Pennington, 2012; Shaywitz,
Shaywitz, Fletcher, & Escobar, 1990). Dyslexia is characterized by difficulties specific to
reading that cannot be explained by hearing difficulty, cognitive deficits, lack of motivation,
or inadequate educational opportunities. Moreover, these difficulties typically persist into
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adulthood (McLoughlin, Leather, & Stringer, 2002). Delineating the underlying mechanisms
that give rise to dyslexia has proven to be an ongoing challenge, as reading is a complex
process. It has been suggested that dyslexia is unlikely to manifest as a singular deficit and
instead may arise from multiple risk factors (Ozernov-Palchik, Yu, Wang, & Gaab, 2016;
Pennington, 2006; van Bergen, van der Leij, & de Jong, 2014). Some of the key deficits
associated with dyslexia include poor phonological awareness (the ability to manipulate
speech sounds within words (Lyon, 2003; Ramus, 2001, 2004; Snowling, 2000)),
weaknesses with phonological working memory (Ramus & Szenkovits, 2008), and difficulty
with rapid automatized naming (Wagner & Torgesen, 1987; Ziegler, Pech-Georgel, Dufau, &
Grainger, 2010). Therefore, the multiple deficit view of dyslexia brings forth consideration
of additional factors that may contribute to this disorder.

Learning to read also relies on nuanced perception and manipulation of speech sounds and
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mapping them to a written, symbolic code (Flax, Realpe-Bonilla, Roesler, Choudhury, &
Benasich, 2009; Nation & Hulme, 1997; Pennington & Lefly, 2001; Snowling, Gallagher, &
Frith, 2003). Accordingly, deficient speech sound perception has been observed in some
individuals with dyslexia (Bogliotti, Serniclaes, Messaoud-Galusi, & Sprenger-Charolles,
2008; Liberman, 1985; Manis et al., 1997; Mody, Studdert-Kennedy, & Brady, 1997;
Vandermosten et al., 2010; Vandermosten et al., 2011). Considerable evidence suggests that
deficits in the perception of basic auditory cues may underlie these speech perception
difficulties (as reviewed in (Hamalainen, Salminen, & Leppanen, 2013)). Therefore, research
has proposed a strong link between early auditory processing, phonological awareness, and
subsequent literacy skills (Tallal, 2004).

Yet, it is puzzling that trained musicians, known to have specialized auditory processing
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skills (Kraus & Chandrasekaran, 2010; Zatorre, Chen, & Penhune, 2007), can also have
persistent dyslexia (Bishop-Liebler, Welch, Huss, Thomson, & Goswami, 2014). This
‘enigma’ of musicians with dyslexia, coined by Weiss and colleagues (2014), calls into
question the extent of a direct link between basic auditory processing and literacy skills in
all individuals with dyslexia. Therefore, investigation of musicians with dyslexia may serve
as one pathway to further investigate the multiple deficit model through the lens of musical

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training, considering multiple risk factors and a complex interplay between perceptual and
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cognitive mechanisms that may give rise to dyslexia (Pennington, 2006).

Although a putative association has been established between basic auditory processing
difficulties early on in development and subsequent dyslexia, the specific nature of this link
remains unclear (Hamalainen et al., 2013). Weaknesses in basic auditory processing have
been reported in individuals with dyslexia for contexts which include discrimination of pitch
and frequency modulation in quiet and in noise (Ahissar, Protopapas, Reid, & Merzenich,
2000; Amitay, Ahissar, & Nelken, 2002; Lorusso, Cantiani, & Molteni, 2014; Tallal &
Piercy, 1973; Wright & Conlon, 2009), and even voice recognition (Perrachione, Del Tufo,
& Gabrieli, 2011). These difficulties have been suggested to manifest from a general
impairment in stimulus-specific prediction, meaning that individuals with dyslexia exhibit
difficulty forming perceptual anchors, which typically allow for increased efficiency and
accuracy with subsequent repetitions of a given stimulus (Ahissar, Lubin, Putter-Katz, &
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Banai, 2006; Oganian & Ahissar, 2012). Within this area of inquiry, one prominent avenue
of investigation is that of non-linguistic temporal processing (Tallal, 2004). Numerous
studies have found that children with language and literacy deficits have difficulties with
discriminating sounds that differ by rapid temporal changes (Tallal & Piercy, 1973, 1974;
Tallal, Stark, & Mellits, 1985). Furthermore, temporal discrimination abilities of infants at
seven months of age have been shown to predict language outcomes at age three (Benasich,
2002), and in turn these abilities in early primary school years have been shown to predict
subsequent literacy skills (Steinbrink, Zimmer, Lachmann, Dirichs, & Kammer, 2014b).

Alternative evidence has indicated that temporal processing difficulties in dyslexia are
specific to perception of the amplitude rise time and slow-rate modulations captured by the
amplitude envelope (Goswami et al., 2002; Lorenzi, Dumont, & Fullgrabe, 2000; Rocheron,
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Lorenzi, Fullgrabe, & Dumont, 2002; Talcott et al., 2000), critical cues for speech
discrimination that convey amplitude changes over time and are known to signify the
rhythmic patterns of speech (Cutler, 1994; Rosen, 1992). Furthermore, temporal processing
deficits in dyslexia have been shown in music-specific contexts, as some children with
dyslexia have demonstrated weaknesses with beat synchronization, rhythm copying, and
rhythmic entrainment (Leong & Goswami, 2014a, 2014b; Overy, Nicolson, Fawcett, &
Clarke, 2003; Thomson, Fryer, Maltby, & Goswami, 2006; Thomson & Goswami, 2008;
Wolff, 2002) as well as discrimination of meter (i.e., the discrimination of beat frequency
and musical accent within rhythmic phrases (Huss, Verney, Fosker, Mead, & Goswami,
2011)).

In addition, basic auditory training has led to improved language and reading abilities in
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children and adults with dyslexia (Gaab, Gabrieli, Deutsch, Tallal, & Temple, 2007; Temple
et al., 2003). Although these studies demonstrate the potential for basic auditory training to
benefit literacy development, further research has importantly revealed that not all
individuals with dyslexia show deficits in auditory processing (Christmann, Lachmann, &
Steinbrink, 2015; Grube, Cooper, Kumar, Kelly, & Griffiths, 2014; Marshall, Snowling, &
Bailey, 2001; Nittrouer, 1999; Ramus, 2003; Rosen, 2003; Steinbrink, Klatte, & Lachmann,
2014a). Indeed, considerable variability has been found in auditory, speech and phonological
processing skills within individuals with dyslexia (Heath, Hogben, & Clark, 1999; Law,

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Vandermosten, Ghesquiere, & Wouters, 2014). In light of these equivocal findings, it is

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particularly intriguing to consider what can be learned about the nature of this relationship
from musicians with dyslexia, as these individuals seem to demonstrate reading difficulties
despite specialized auditory skills in the musical domain.

Musical training has been put forth as a promising tool to promote auditory specialization
and therefore support literacy skill development (Rolka & Silverman, 2015; Tallal & Gaab,
2006). Individuals with musical training have demonstrated heightened discrimination skills
over non-musicians for several components of auditory processing, including spectral
features such as pitch (Amir, Amir, & Kishon-Rabin, 2003; Besson, Schon, Moreno, Santos,
& Magne, 2007; Carey et al., 2015; Kishon-Rabin, Amir, Vexler, & Zaltz, 2001; Koelsch,
Schroger, & Tervaniemi, 1999; Magne, Schon, & Besson, 2006; Micheyl, Delhommeau,
Perrot, & Oxenham, 2006; Spiegel & Watson, 1984), and temporal features such as elements
of timing (Cicchini, Arrighi, Cecchetti, Giusti, & Burr, 2012; Ehrle & Samson, 2005; Gaab
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et al., 2005; Rammsayer & Altenmüller, 2006).

These pitch and timing cues are not only necessary for music but are also critical to speech
processing, which demands precise perception of the formant frequencies that characterize
vowels and consonants within a specific, rapid temporal framework (Stevens, 1980).
Overlapping spectral and temporal features across music and speech perception suggest that
the auditory specialization achieved through intensive musical training may also be
associated with advantageous speech processing abilities (Chandrasekaran & Kraus, 2010;
Chobert, Francois, Velay, & Besson, 2012; Parbery-Clark, Skoe, & Kraus, 2009; Patel, 2011,
2012). Accordingly, trained musicians have shown superior detection of spectral features
within speech compared to non-musicians (Deguchi et al., 2012; Schon, Magne, & Besson,
2004; Thompson, Schellenberg, & Husain, 2003), as well as heightened perception of
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temporal speech-specific features for properties such as segmental structure (Francois,

Chobert, Besson, & Schon, 2012; Moreno et al., 2009) and the amplitude envelope (Zuk et
al., 2013b). In addition, specialized electroencephalographic and auditory brainstem
responses during sound discrimination have been reported in musicians over non-musicians
for music and speech stimuli characterized by differences in frequency, duration, and
intensity (Jentschke, Koelsch, & Friederici, 2005; Moreno & Besson, 2006; Tervaniemi et
al., 2009; Weiss & Bidelman, 2015; Wong, Skoe, Russo, Dees, & Kraus, 2007).
Furthermore, longitudinal studies have shown changes in these neural responses to speech
following musical intervention in childhood (Habibi, Cahn, Damasio, & Damasio, 2016;
Kraus, Hornickel, Strait, Slater, & Thompson, 2014a; Kraus et al., 2014b, 2014c). These
specialized neural responses to speech in adults with musical training have also been shown
to significantly relate to the total amount and intensity of musical training (Musacchia,
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Strait, & Kraus, 2008), suggesting that long-term dedication to musical training that involves
intense practice routines may facilitate distinct mechanisms for speech processing. Thus, a
growing body of evidence supports the notion that musical training may serve as an effective
outlet to advance basic auditory and speech-specific processing skills, which in turn may
positively impact reading skills (Tallal & Gaab, 2006).

Musicianship has furthermore been directly linked with early and developing language and
literacy skills (Fisher & McDonald, 2001; Moritz, Yampolksy, Papadelis, Thomson, & Wolf,

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2012; Williams, Barrett, Welch, Abad, & Broughton, 2015). Musical training as well as
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musical aptitude (as indicated by music perception tasks) have shown positive associations
with phonological abilities such as rhyming, blending, sound isolation, and segmentation
(Anvari, Trainor, Woodside, & Levy, 2002; Forgeard, Schlaug, Norton, Rosam, & Iyengar,
2008; Loui, Kroog, Zuk, Winner, & Schlaug, 2011; Standley & Hughes, 1997; Zuk,
Andrade, Andrade, Gardiner, & Gaab, 2013a). Moreover, music-based intervention has
resulted in improved phonological processing and speech segmentation in typically
developing school-age children as well as struggling readers (Bhide, Power, & Goswami,
2013; Dege & Schwarzer, 2011; Hurwitz, Wolff, Bortnick, & Kokas, 1975; Moreno et al.,
2009; Overy, 2003; Przbylski et al., 2013; Santos, Joly-Pottuz, Moreno, Habib, & Besson,
2007; Thomson, Leong, & Goswami, 2013).

Positive relationships have also been found between musical skill and various reading
abilities including reading speed and accuracy (Barwick, Valentine, West, & Wilding, 1989;
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Corrigall & Trainor, 2011; Douglas & Willatts, 1994; Gardiner, Fox, Knowles, & Jeffrey,
1996; Goswami, Huss, Mead, Fosker, & Verney, 2012; Hurwitz et al., 1975; Lamb &
Gregory, 1993; Register, Darrow, Standley, & Swedberg, 2007; Standley & Hughes, 1997;
Strait, Hornickel, & Kraus, 2011; Zuk et al., 2013a). In addition, music-based interventions
in children with dyslexia have shown improvements in phonological awareness (Atterbury,
1985; Farmer, Kittner, Rae, Bartko, & Regier, 1995; Flaugnacco et al., 2015; Habib et al.,
2016; Overy, 2003; Santos et al., 2007; Thomson et al., 2013), spelling (Atterbury, 1985;
Farmer et al., 1995; Overy, 2003; Santos et al., 2007), and reading skills (Flaugnacco et al.,
2015; Habib et al., 2016; Thomson et al., 2013). However, the extent of these gains warrants
further investigation as the only study to include multiple control groups found no
significant differences in improvements between children who received six weeks of
computer-based rhythm training and others who received a traditional language-based
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intervention, even compared to controls who did not engage in any specific training
(Thomson et al., 2013). Thus, although a connection between music and literacy is evident,
the nature of this relationship has yet to be sufficiently specified.

Despite the collective evidence linking auditory processing deficits with reading difficulties
and the potential to remediate these skills through musical training, there are numerous cases
of individuals with dyslexia with persistent difficulties who study classical music at the
highest levels and become professional musicians (Bishop-Liebler et al., 2014). Thus, this
population raises significant questions as to whether individuals with dyslexia who have
received extensive musical training evidence specific advantages in auditory processing
relative to those with dyslexia who have not had musical training. If so, questions remain as
to why these individuals still develop (persistent) literacy difficulties, and to what extent
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their putative advantages in auditory processing reflect a direct influence of musical training
or rather an early propensity for specialized auditory processing.

To date, only two studies have investigated auditory processing skills in musicians with
dyslexia. The first study to examine this compared musicians with dyslexia to typical
musicians on a wide variety of auditory processing abilities, including non-linguistic spectral
and temporal discrimination, speech sound perception in noise, synchronous finger tapping,
and auditory verbal and non-verbal working memory (Weiss, Granot, & Ahissar, 2014).

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Musicians with dyslexia performed comparably with typical musicians on all of the basic
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auditory processing and finger tapping tasks. Yet, musicians with dyslexia demonstrated
poor auditory working memory performance relative to typical musicians for phonological
and musical stimuli, thus contributing to the body of literature that attributes poor working
memory to be a critical deficit underlying dyslexia (de Jong, 1998; Wagner & Torgesen,
1987).

The second investigation of musicians with dyslexia to date also identified no significant
differences in auditory processing skills between this population and typical musicians, as
measured by discrimination tasks of frequency, intensity, the amplitude rise time conveyed
by the amplitude envelope, and rhythm perception (Bishop-Liebler et al., 2014).
Additionally, musicians with dyslexia performed better than non-musicians with dyslexia on
the majority of these measures, but did not demonstrate a significant advantage on amplitude
rise time perception or the duration discrimination task. Thus, extant findings suggest that
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musicians with dyslexia exhibit specialized auditory processing skills similar to typical
musicians for certain auditory constituents but not all, which is conceivable since the
difficulties associated with dyslexia typically continue into adulthood (McLoughlin et al.,
2002). Yet, it remains unclear whether musicians with dyslexia also exhibit specific deficits
with speech processing as previously shown in individuals with dyslexia (Bogliotti et al.,
2008; Liberman, 1985; Manis et al., 1997; Mody et al., 1997; Vandermosten et al., 2010;
Vandermosten et al., 2011).

As reviewed above, investigations of musicians with dyslexia have primarily focused on

specific components of auditory processing, and characterization of speech-specific
processing abilities remain largely unspecified. Further investigation of the abilities of
musicians with dyslexia for non-speech and speech-specific perception tasks has the
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potential to provide further insight on the extent of the ‘musician advantage’ this unique
population possesses, and the auditory expertise that may be observable despite persistent
reading difficulties. As such, the lack of differences in auditory processing abilities in
musicians with dyslexia relative to typical musicians brings forth a question of whether these
‘auditory advantages’ extend to speech-specific contexts. Research evidence has yet to
uncover whether the superior speech-specific processing abilities shown in typical musicians
in the spectral (Deguchi et al., 2012; Magne et al., 2006; Schon et al., 2004; Thompson et
al., 2003) and temporal (Francois et al., 2012; Moreno et al., 2009; Zuk et al., 2013b)
domains may also be evident within musicians with dyslexia. Furthermore, prior evidence
has suggested weaknesses in auditory working memory in musicians with dyslexia
compared to musicians without (Weiss et al., 2014). Accordingly, it is unclear whether
musicians with dyslexia may be characterized by working memory deficits in general, or
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whether the auditory expertise afforded by musical training may be associated with some
advantages in processing auditory information even when taxing the working memory
system. Auditory sequencing is particularly of interest since attending to and reproducing
auditory sequences is one of the primary auditory skills developed through musical training
(Carey et al., 2015; Loui, Wessel, & Hudson Kam, 2010; Rohrmeier, Rebuschat, & Cross,
2011; van Zuijen, Sussman, Winkler, Naatanen, & Tervaniemi, 2005). Moreover, auditory
sequencing has also been shown to be a critical building block for language (Tallal & Gaab,

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2006; Tallal et al., 1985). To date, auditory sequencing abilities have yet to be investigated in
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musicians with dyslexia.

The present study will advance the extant indicators of auditory processing abilities in
musicians with dyslexia through a battery of tasks measuring non-speech and speech-
specific auditory processing. As with previous investigations of musicians with dyslexia, this
study precludes determination of whether the putative specialization in musicians with
dyslexia may be the direct result of long-term musical training, or instead a predisposition
for musical achievement. Even so, the present study will further characterize auditory
processing abilities in musicians with dyslexia through tasks that have not yet been utilized
to assess musicians with dyslexia. These tasks include (i) an auditory processing task
measuring tone sequencing skills and (ii) speech-specific perceptual tasks that have been
previously employed to explore processing abilities in musicians (Zuk et al., 2013b).
Specifically, these speech tasks measure discrimination thresholds of synthetic syllable
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continua that vary in spectral (frequency) and temporal (amplitude envelope and voice onset
time) features. Performance in musicians with dyslexia will be directly compared with that
of typical musicians, as well as non-musicians with dyslexia.

Consequently, we identified two hypotheses. First, we expected to find no significant

differences in accuracy on the tone-sequencing task between musicians with dyslexia and
typical musicians, consistent with prior findings of no significant differences between these
groups for auditory processing tasks (Bishop-Liebler et al., 2014; Weiss et al., 2014). By
comparison, nonmusicians with dyslexia were hypothesized to show significantly poorer
performance relative to the groups with musical training. Yet, we also anticipated that
musicians with dyslexia may reveal slower reaction times than typical musicians for the tone
sequencing task, based on the evidence of weaknesses in speed of processing of individuals
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with dyslexia (Breznitz & Misra, 2003; Catts, Gillispie, Leonard, Kail, & Miller, 2002). The
second hypothesis was for the speech-specific tasks that a certain degree of specialization
would be evident in musicians with dyslexia, since we expected that the reported advantages
in auditory processing in this group would extend to speech-specific contexts as well. Yet,
musicians with dyslexia were hypothesized to perform more similarly to nonmusicians with
dyslexia for the discrimination of syllable continua requiring distinction of temporal features
of speech, considering the significant evidence for deficits in dyslexia in discerning temporal
information (Goswami et al., 2002; Lorenzi et al., 2000; Rocheron et al., 2002; Talcott et al.,
2000; Vandermosten et al., 2010; Vandermosten et al., 2011). In particular, these groups
were expected to perform comparably for discrimination of syllables that vary in amplitude
envelope cues with superior performance in typical musicians, based on prior evidence of
similar performance between musicians and nonmusicians with dyslexia for detection of
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amplitude rise time cues in a non-speech context (Bishop-Liebler et al., 2014). Thus, the
present investigation sought to characterize specialized auditory sequencing and speech
processing skills and identify whether persistent weaknesses were evident in musically
trained individuals with dyslexia. Furthermore, the present study sought to uncover the
extent of an association between musical training and specific aspects of auditory processing
critical to early literacy development. Taken together, these aims may provide implications
for the potential of musical training to benefit auditory and literacy skill development in
individuals with dyslexia.

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Methods
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Participant Demographics
Fifty-two healthy, monolingual, native British English-speaking adults were included in the
present study (17 male, 35 female, ages 18–36 years with mean age: 20.89 yrs, SD: 2.82).
Three groups classified participants as follows: typical musicians (TYPMUS; n = 17, 14
female, mean age: 21 yrs, STD: 1.73), musicians with dyslexia (DYSMUS; n = 19, 8 female,
mean age: 20.68 yrs, STD: 2.29), and nonmusicians with dyslexia (DYSNonMUS; n = 16,
13 female, mean age: 21.06 yrs, STD: 4.17). All participants were UK university students or
recent graduates, recruited through student service departments, music departments, or
academic departmental postings at their institutions. All participants with dyslexia had
previously received a formal diagnosis from an educational psychologist or qualified
specialist. Formal diagnosis of dyslexia among participants adhered to UK definitions of
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dyslexia and evaluation protocols, as described by the Department of Education and Schools
Guidelines (DfES, 2005). The age of dyslexia diagnosis in these participants ranged from
childhood to adulthood (mean age: 17.63 years, SD: 3.85, with no significant differences in
musician vs. nonmusician groups, though three participants did not provide this
information). Musicians (TYPMUS and DYSMUS) in this study were defined as either
being enrolled in or having obtained a music performance degree specializing in classical or
jazz music (see Table 1 for details of musical training). On average, musicians in both
groups began studying music at a mean age of seven years and had completed approximately
thirteen years of musical training (as shown in Table 1 by group). DYSNonMUS had no
prior musical training outside of the requirements of the general music curriculum in school.
Participants were confirmed to have no neurological abnormalities, hearing impairments, nor
additional neuropsychological or developmental diagnoses. The three groups showed no
significant differences in age or nonverbal IQ (see Table 3 for an overview of IQ scores).
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Ethical approvals for this collaborative study were granted by Boston Children’s Hospital,
the Institute of Education at University College London, and the Edinburgh College of Art at
the University of Edinburgh. All participants provided written informed consent.

Literacy Measures
Participants were characterized by a battery of standardized assessments that evaluated
language and literacy abilities. Phonological awareness skills were assessed through the
Elision and Blending subtests of the Comprehensive Test of Phonological Processing
(CTOPP; (Wagner, Torgesen, & Rashotte, 1999)). Rapid automatized naming skills were
measured through the Rapid Digit Naming and Rapid Letter Naming subtests of the CTOPP.
Composite standard scores for phonological awareness and rapid naming were calculated.
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Phonological working memory was measured through composite score on Digit Backwards
and Digit Forward subtests of the Digit Memory Test (Turner & Ridsdale, 2004). Word
reading and spelling were evaluated through the Wide Range Achievement Test (WRAT;
(Wilkinson, 1993; Wilkinson & Robertson, 2006). Rapid (timed) single word reading was
assessed through subtests of the Test of Word Reading Efficiency (TOWRE; (Torgesen,
Wagner, & Rashotte, 1999)): Sight Word Efficiency (timed single-word reading) and
Phonemic Decoding (timed decoding of non-words). Verbal and Nonverbal IQ were
determined by the mean standard score of the two verbal subtests (Verbal Analogies/

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Similarities and Vocabulary) and two nonverbal subtests (Diamonds and Matrices) of the
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Wide Range Intelligence Test (WRIT; (Glutting, Adams, & Sheslow, 2000)) or equivalents
from the Wechsler Adult Intelligence Scale (WAIS-IIIUK; Wechsler, 1998). The WAIS and
WRIT are strongly correlated and are therefore reported interchangeably (DfES, 2005).
Inclusion criteria for participation in this study required a standard score of no less than one
standard deviation below the mean (>85) on any of the IQ subtests. All participants in the
TYPMUS group achieved scores within and above the average range on all measures of
phonological processing, reading, and spelling (i.e., standard scores > 90).

In addition to a formal diagnosis of dyslexia, inclusion criteria were set based on reading
and spelling achievement to validate the accuracy of self-reported diagnosis and ensure that
a representative sample of individuals with dyslexia has been included in the DYSMUS and
DYSNonMUS groups relative to the general population of adults with dyslexia (McLoughlin
et al., 2002). Specifically, participants with dyslexia met the criteria for this study if they
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obtained a standardized score below 90 on at least one of the TOWRE subtests (Sight Word
Efficiency or Phonemic Decoding). Accordingly, additional participants with dyslexia were
excluded from analysis due to high scores on reading measures. In the case that participants
with dyslexia provided the research team with a full diagnostic report that had been
conducted within four years of study participation, standardized scores from measures that
would have been re-administered in the present research study were taken from the
diagnostic report. Thus, 52 participants as described above were included in the present
analysis (TYPMUS n = 17; DYSMUS n = 19; DYSNonMUS n = 16).

Tone Sequencing Task

Tone sequences comprised of two complex tones with fundamental frequencies of 100 (low
pitch) and 300 Hz (high pitch) in a modification of Tallal’s Repetition Test (Tallal & Piercy,
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1973). Both tones included eight harmonics with a six-decibel drop off, were equalized for
power using a root-mean-squared formula, and had durations of 50, 75 and 125 ms. Tones
were presented with different inter-stimulus intervals (ISIs) of 5, 10, 20, 40, 80 or 160 ms.
For any given trial, all tones were of the same duration (50, 75, or 125 ms) and the ISI was
constant (either 5, 10, 20, 40, 80 or 160 ms).

The tone sequencing task required participants to make a motor response (via button press)
to indicate the order in which they heard the low- and high-pitched tones in each trial. The
task began with training the motor response to each tone presented separately before
progressing to replication of tone sequences of increasing length (two-, three- and four-tone
sequences). Accuracy was measured by correct indication of low- and high-pitched tones in
the same order as presented in each trial (any error within the sequence resulted in a score of
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zero for the trial). Reaction time for each trial was also acquired in all participants.

Syllable Task Stimuli

The syllable task comprised of the following three synthetic speech syllable continua: /
bɑ/-/dɑ/ (spectral change within formant transition), /bɑ/-/wɑ/ (duration change of formant
transition/amplitude envelope), and /ɡɑ/-/kɑ/ (change in Voice Onset Time; see Figure 1).
The /bɑ/-/dɑ/ contrast was defined by direct changes in the onset of the second formant with

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a constant duration of the formant transition (40 ms) for all stimuli in the continuum. The /
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bɑ/-/wɑ/ continuum involved manipulation of the amplitude envelope (as described in (Zuk
et al., 2013b)), while the /ɡɑ/-/kɑ/ continuum was created by altering the Voice Onset Time
(VOT), both primarily involving a temporal change. Syllable stimuli were created through a
Klatt-based synthesizer (Klatt, 1980). All syllables were 250 ms in duration and had a
fundamental frequency (F0) of 120 Hz, which dropped to 90 Hz through the duration of the
syllable. The specifications of the acoustic parameters used to synthesize the three continua
were the following:

/bɑ/-/dɑ/ continuum: The onset value of the second formant for the /bɑ/-/dɑ/ continuum
varied from 800 to 1600 Hz, (/bɑ/ and /dɑ/, respectively), in 32 Hz steps producing 26
syllables spanning a spectral continuum between /bɑ/ and /dɑ/. The starting frequencies for
the formant transitions of the /bɑ/-/dɑ/ continuum were: F1 = 420 Hz, F2: varying from 800
to 1600 Hz, F3=2500 Hz, F4=3250 Hz, and F5=3700 Hz. The transition was 40 ms, at
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which point the formant frequency (F) and bandwidth (BW) values were: F1 = 800 Hz, BW1
= 90; F2=1200 Hz, BW2=110; F3=2500 Hz, BW3=90; F4=3250 Hz, BW4=400; F5=3700
Hz, BW5=500. At 180 ms, the formant frequency changes were: F1 = 750 Hz and the
voicing was ramped down to zero for the remaining duration.

/bɑ/-/wɑ/ continuum: The duration of the transition varied from 25 to 97 ms (/bɑ/ and /wɑ/,
respectively), in steps increasing by three milliseconds each, producing 25 syllables along
this continuum. The frequency and bandwidth specifications were identical to the /bɑ/ used
in the /bɑ/-/dɑ/ continuum (see above) except F2 remained 800 Hz and the transition
duration varied from 25 to 106 ms. This continuum has also been characterized by changes
in the amplitude rise time duration, as previously analyzed by Zuk, Ozernov-Palchik and
colleagues (2013).
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/ɡɑ/-/kɑ/ continuum: The Voice Onset Time (VOT) for each syllable in the /ɡɑ/-/kɑ/
spectrum ranged from 10 to 60 ms, (/ɡɑ/ and / kɑ/, respectively) in two millisecond steps
producing 26 syllables along this continuum. The starting frequencies for the formant
transitions were: F1 = 300 Hz, F2 = 1625 Hz, F3 = 2000 Hz, F4 = 3250 Hz, and F5 = 3700
Hz. The formant frequency and bandwidth values at the beginning of the vowel were:
F1=700 Hz, BW1=90; F2=1200 Hz, BW2=90; F3=2300 Hz, BW3=130; F4=3300 Hz,
BW4=400; F5=3700Hz, BW5=500. At 180ms, the formant frequency changes were: F1 =
750 Hz, F2 = 1000 Hz, F3 = 2300 Hz and the voicing was ramped down to zero for the
remaining duration.

Participants were presented with a pair of syllables (one after the other with an inter-
stimulus interval of 750 ms) and asked to indicate whether the two syllables sounded the
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same or different via button press. Each pair contained a fixed reference syllable (/bɑ/, /bɑ/,
or /ɡɑ/ depending on the continuum) and a test syllable. The presentation order of the
reference and test syllable was randomized throughout the task. The task progressed through
trials in accordance with the three-down one-up adaptive staircase method
(Lakshminarayanan & Tallal, 2007). At the onset of the task, the test syllable was at the
opposite end of the continuum from the reference syllable; that is, trials always began with
the most easily discriminable stimulus pair from the continuum. Specifically, the

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discrimination limen of the first stimulus pair in the /bɑ/-/dɑ/ continuum corresponded to the
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syllable with a second formant frequency of 800 Hz (/bɑ/) and 1600 Hz (/dɑ/), marking the
extremes of the continuum. Accordingly, the discrimination limen of the first pair for the /
bɑ/-/wɑ/ continuum was 97 ms and for the /ɡɑ/-/kɑ/ continuum, 60 ms.

After three consecutive correct responses to the first syllable pair, the discrimination limen
decreased by two steps and the trials progressed accordingly. For each incorrect response,
the discrimination limen increased by one step and an easier stimulus pair in the continuum
was presented until seven reversals in the direction of progression of trials were achieved.
Catch trials containing pairs of identical syllables were presented every 5–10 trials (for
which all participants performed at 100%). Each assessment was terminated after seven
reversals or five consecutive incorrect responses to the initial, most easily distinguishable
pair. The discrimination thresholds for each of the stimulus continua were determined by the
arithmetic mean of the discrimination limen corresponding to the last four reversals. The
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original threshold value was measured in Hz for /bɑ/-/dɑ/ and in ms for /bɑ/-/wɑ/ and /
ɡɑ/-/kɑ/. Prior to commencing the task, participants completed a practice session of five
syllable pairs to familiarize themselves with the stimuli and ensure they understood the
instructions. In order to allow for direct comparison between the three syllable continua, the
discrimination thresholds in Hz and ms were transformed into a Relative Threshold Index
(RTI) ranging from zero to one (as previously described in (Zuk et al., 2013b); see Table 2
for conversion formulas).

Specifically, the RTI was the value obtained by subtracting the reference syllable value
(for /bɑ/-/dɑ/ 800 Hz, for /bɑ/-/wɑ/ 25 ms and for /ɡɑ/-/kɑ/ 10 ms) from the obtained
discrimination threshold. This number was then divided by the maximum range for each
acoustic continuum (for /bɑ/-/dɑ/ 800 Hz, for /bɑ/-/wɑ/ 97 ms and for /ɡɑ/-/kɑ/ 60 ms) and
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subtracted from 1 (see Table 2). Thus, a higher RTI indicates better discrimination. For
example, a discrimination threshold of 1400 Hz for the /bɑ/-/dɑ/ continuum would equate to
an RTI of 0.25, while a discrimination threshold of 1000 Hz would be designated by an RTI
of 0.75.

General Procedure and Analysis

For both experiments, participants were seated comfortably in a quiet testing room with a
PC computer running ePrime (Psychology Software Tools, 2002). Stimuli were transmitted
through Panasonic and Beyer stereo headphones. Comparisons between groups on language
and literacy measures, the tone sequence task and syllable task performance were evaluated
through one-way ANOVAs, repeated measures ANOVAs and post-hoc Games-Howell
calculations. A priori, a sample size of at least 50 participants was estimated to be necessary
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to achieve at least 80% power with a large effect size (i.e., ≥0.4 based on standard effect size
conventions for ANOVAs (Cohen, 1988) and the effect sizes of related previous studies), and
an alpha-level threshold of 0.05 for ANOVA analyses with three groups and post-hoc group
comparisons. Therefore, our sample size of 52 participants was deemed suitable to estimate
group differences with sufficient power. In addition, post-hoc effect sizes were calculated to
ensure that the expected effect sizes have been achieved. Specifically, effect sizes were
estimated based on standard formulas for eta squared (η2) from ANOVA analyses

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(Rosenthal & Rosnow, 1985, 1991), and the effect size index d as defined by Cohen for post-
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hoc comparisons of mean differences between groups (Cohen, 1969). Participants included
in the present analysis completed administration of all three syllable contrasts; participants
who were only administered one-to-two syllable contrasts due to time constraints were not
included in the present analysis.

Results
Literacy Demographics
One-way ANOVAs investigating group differences on literacy-related measures revealed
significant group differences for all measures of phonological processing, reading, and
spelling (all significant at p < 0.001; see Table 3 for F-values corresponding to each specific
measure). Post-hoc Games-Howell tests for unequal variances showed that TYPMUS
performed significantly better than both DYSMUS and DYSNonMUS on all measures (all
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significant at p < 0.001; see Table 3 for an overview) other than Nonverbal IQ, as expected
given our inclusion criteria. Post-hoc direct comparison between DYSMUS and
DYSNonMUS demonstrated that DYSMUS achieved significantly higher scores on the
Sight Word Efficiency subtest than DYSNonMUS with a large effect size (Mean Difference
(MD) = 6.984, Standard Error (SE) = 2.4, p = 0.017, d = 0.981), and these groups otherwise
did not significantly differ on any other measures.

Tone Sequencing Task

For accuracy on the tone sequencing task, a repeated measures ANOVA (with group as the
between-participant factor and the number of tones in the sequence as the within-participant
factor) was implemented. First, inclusion criteria were established to ensure appropriate
completion of the task, and that the participants had understood the task correctly. All
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participants achieved nearly 100% accuracy on all the single tone trials, indicating that they
had no difficulty discriminating the pitch of the tones and understood the required key press
response. Five DYSNonMUS did not complete this task due to time constraints during data
collection (i.e., this task thus included the following number of participants in each group:
TYPMUS: n = 16; DYSMUS: n = 19; DYSNonMUS: n = 11).

An ANOVA with repeated measures revealed significant differences in accuracy between

groups with a large effect size (F(2,43) = 65.821, p < 0.001, η2 = 0.754). Post-hoc Games-
Howell tests revealed that TYPMUS and DYSMUS performed comparably (MD = 0.04, SE
= 0.026, p = 0.268, d = 0.454), and both musician groups were significantly more accurate
than DYSNonMUS for all tone sequences (all comparisons resulting in p < 0.001, d > 0.4).
As shown in Figure 2, participants in all groups were significantly less accurate as the
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number of tones in each sequence increased (F(2,42) = 136.762, p < 0.001, η2 = 0.867).
Within each tone sequence, differences between the TYPMUS and DYSMUS groups
relative to DYSNonMUS decreased as the inter-stimulus interval (ISI) increased for the two-
tone and three-tone sequences, however, performance steadily increased in all three groups
as ISI increased for the four-tone sequence. Despite distinct patterns of differences in
performance between groups as tone sequence and ISI increased, one-way ANOVAs and
post-hoc Games-Howell tests revealed significant differences between DYSNonMUS and

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the TYPMUS and DYSMUS groups for all ISIs at each tone sequence (all comparisons
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resulting in p < 0.001, d > 0.4). Ceiling effects were observed within both TYPMUS and
DYSMUS for all ISIs of the two-tone sequences and at an ISI of 80 and 160 ms for the
three-tone sequence (see Figure 2). All groups performed above chance on average for each
tone sequence (as indicated in Figure 2).

Reaction time on the tone sequence task was also found to significantly differ between
groups, as revealed by ANOVA with a medium effect size (F(2,43) = 70.182, p < 0.001, η2 =
0.336). Post-hoc Games-Howell tests revealed that TYPMUS and DYSMUS showed no
significant differences in reaction time (MD = 28.489 msec, SE = 21.862 msec, p = 0.395, d
= 0.08), albeit with a small effect size, whereas both TYPMUS and DYSMUS had
significantly faster reaction times than DYSNonMUS on all tone sequences with large effect
sizes (all comparisons resulting in p < 0.001, d > 0.8). As expected, reaction time
significantly increased as the number of tones in each sequence increased for all three
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groups with a large effect size (F(2,42) = 661.346, p < 0.001, η2 = 0.827).

Syllable Task
Relative Threshold Index (RTI) outcomes for the three syllable continua were evaluated
through ANOVA with repeated measures. Despite sufficient performance on practice trials
immediately prior to commencing the task, ten participants with dyslexia (five DYSMUS,
five DYSNonMUS) were unable to move forward on certain syllable continua beyond the
first discrimination limen. After five consecutive incorrect responses to the initial, most
easily distinguishable pair the task terminated. Consequently, these participants were
assigned the first step in the continuum as their discrimination threshold and given an RTI
score of zero for that particular syllable continuum. Instances of termination occurred in ten
participants within specific syllable continua as follows: three for the /bɑ/-/dɑ/ continuum,
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two for /bɑ/-/wɑ/, and eight for the /ɡɑ/-/kɑ/ continuum (two of which also did not achieve
beyond the first pair for /bɑ/-/dɑ/, and one who was also terminated on /ba/-/wa/).

A series of ANOVAs confirmed significant between-subject group differences on all three of

the syllable continua (see Table 4 for an overview). Direct group comparisons with post-hoc
Games-Howell tests confirmed that for the /bɑ/-/dɑ/ continuum, RTI scores did not
significantly differ between TYPMUS and DYSMUS (MD = 0.024, SE = 0.024, p = 0.595, d
= 0.33), and these two groups demonstrated superior discrimination thresholds than
DYSNonMUS with large effect sizes (comparisons resulted in p < 0.05, d > 0.8). Similar
group differences were found for the /ɡɑ/-/kɑ/ continuum varying in Voice Onset Time
(VOT), in which post-hoc evaluation revealed no significant differences in discrimination
thresholds between TYPMUS and DYSMUS (MD = 0.062, SE = 0.076, p = 0.699, d =
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0.27). Although a relatively small effect size was found when comparing TYPMUS and
DYSMUS, large effect sizes resulted from other group comparisons, in which TYPMUS
demonstrated significantly better discrimination thresholds than DYSNonMUS for /ɡɑ/-/kɑ/
(MD = 0.229, SE = 0.07, p = 0.007, d = 1.14), and differences between DYSMUS and
DYSNonMUS trended towards significance (MD = 0.167, SE = 0.07, p = 0.056, d = 0.81).
Lastly, TYPMUS only showed heightened discrimination thresholds over DYSMUS for the
continuum varying in amplitude envelope, /bɑ/-/wɑ/ (MD = 0.219, SE = 0.07, p = 0.013, d =

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1.03), who in turn did not significantly differ from DYSNonMUS (MD = 0.08, SE = 0.093, p
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= 0.699, d = 0.3). An overview of group differences on each of these syllable continua is

provided in Figure 3.

Discussion
The present investigation of auditory sequencing and speech processing abilities in adult
musicians with dyslexia who evidenced persistent literacy difficulties has revealed a distinct
profile of abilities relative to both nonmusicians with dyslexia and typical musicians. As
expected, typical musicians performed significantly better than musicians and nonmusicians
with dyslexia on all literacy-related tasks and exhibited significantly better verbal
intelligence scores, though all groups demonstrated above average mean scores on this
measure. Musicians and nonmusicians with dyslexia did not significantly differ on these
standardized measures except for the sight word reading efficiency subtest, in which
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musicians with dyslexia showed better performance than nonmusicians with dyslexia.
Characterization of tone sequencing abilities revealed, as expected, that the accuracy of
musicians with dyslexia did not significantly differ from typical musicians, whereas
nonmusicians with dyslexia performed significantly more poorly than both musician groups
for all tone sequences and inter-stimulus intervals. As for the speech-specific perceptual
tasks, which measured the discrimination thresholds of synthetic speech syllable continua,
musicians with dyslexia did not differ from typical musicians for discrimination of two out
of three acoustic properties isolated within the syllable continua. Specifically, musicians
with dyslexia achieved discrimination thresholds that did not significantly differ from those
of typical musicians on measures distinguishing spectral and VOT cues within syllables.
Both musician groups also achieved significantly better discrimination thresholds than
nonmusicians with dyslexia for these tasks. Yet, musicians with dyslexia did not differ from
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nonmusicians with dyslexia for discrimination of the /bɑ/-/wɑ/ contrast, which specifically
evaluated the discrimination of amplitude envelope cues. With predominantly medium to
large effect sizes corresponding to the present findings, it is evident that musicians with
dyslexia showed advantages in auditory sequencing compared to nonmusicians with
dyslexia. Musicians with dyslexia also showed refined speech discrimination abilities for
two critical acoustic features, but demonstrated relative difficulties with discrimination of
syllables varying by amplitude envelope.

The present findings suggest that musical training is associated with specialized auditory
sequencing abilities in those with dyslexia, despite findings of poor auditory working
memory performance in musicians with dyslexia. These results support our hypothesis that
despite reading difficulties, musicians with dyslexia exhibit tone sequencing skills similar to
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those that have been shown to be developed and mastered in typical musicians (Carey et al.,
2015; Loui et al., 2010; Rohrmeier et al., 2011; van Zuijen et al., 2005). These refined tone
sequencing abilities in musicians with dyslexia are also in line with prior findings of
auditory perception skills that did not significantly differ between musicians with dyslexia
and typical musicians for non-linguistic auditory tasks (Bishop-Liebler et al., 2014; Weiss et
al., 2014). Conversely, nonmusicians with dyslexia demonstrated poor tone sequencing skills
relative to musician groups, which is in line with previous reports of non-linguistic auditory

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processing deficits in individuals with dyslexia (Christmann et al., 2015; Grube et al., 2014;
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Marshall et al., 2001; Nittrouer, 1999; Ramus, 2003; Rosen, 2003; Steinbrink et al., 2014a).

Although promising findings emerge from the tone-sequencing task, two significant
considerations are important to note. First, a ceiling effect was observed in both musician
groups for the two- and three-tone sequences, which makes it unclear whether a more
complex non-speech task may further distinguish typical musicians relative to musicians
with dyslexia. The second consideration concerns the validity of this task in measuring
specifically auditory processing, and to what extent working memory may be taxed as the
number of tones in each sequence increases. This should be taken into account, as prior
studies have found auditory processing deficits to be concomitant with working memory
difficulties (Ahissar et al., 2000; Banai & Ahissar, 2004). In the present sample, working
memory, as measured behaviorally by the digit span test, revealed that typical musicians
were superior to both groups with dyslexia, and musicians with dyslexia did not significantly
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differ from nonmusicians with dyslexia in digit span achievement, consistent with prior
findings (Weiss et al., 2014). Therefore, despite working memory weaknesses relative to
typical musicians, musicians with dyslexia demonstrated strengths in the present tone
sequencing task. This finding is in line with the notion put forth by Weiss and colleagues
(2014) that musicians with dyslexia seem to demonstrate divergent auditory processing
skills, in which perception of auditory constituents in general present as a relative strength
with significant weaknesses in auditory working memory.

As for speech-specific perceptual abilities in musicians with dyslexia, syllable

discrimination thresholds significantly differed between musicians with dyslexia from both
typical musicians and nonmusicians with dyslexia. For these tasks, it was hypothesized that
a certain degree of specialization would be evident in musicians with dyslexia due to the
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considerable evidence that musical training is associated with refined speech processing
abilities (Chandrasekaran & Kraus, 2010; Chobert et al., 2012; Parbery-Clark et al., 2009;
Patel, 2012; Tallal & Gaab, 2006). In support of this hypothesis, syllable discrimination did
not significantly differ between musicians with dyslexia and typical musicians for
discrimination of spectral cues (/bɑ/-/dɑ/) and a temporal acoustic cue, voice onset time
(VOT, as indicated by /ɡɑ/-/kɑ/). These findings align well with the growing evidence that
trained musicians demonstrate specialized speech perception abilities in both spectral
(Deguchi et al., 2012; Magne et al., 2006; Schon et al., 2004; Thompson et al., 2003) and
temporal (Francois et al., 2012; Moreno et al., 2009; Zuk et al., 2013b) domains.
Furthermore, these findings are in line with longitudinal studies that have shown enhanced
neural responses in the auditory brainstem to speech stimuli following musical training
(Kraus et al., 2014a; Kraus et al., 2014b, 2014c), suggesting that musical training may
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facilitate distinct mechanisms for speech processing.

For distinction of the /bɑ/-/dɑ/ and /ɡɑ/-/kɑ/ contrasts, both musician groups were superior
to nonmusicians with dyslexia. Poorer performance in nonmusicians with dyslexia was
expected, since deficient speech sound representations have been found in individuals with
dyslexia relative to controls for similar tasks involving discrimination of synthetic speech
stimuli that varied in spectral and temporal features (Bogliotti et al., 2008; Manis et al.,
1997). The present findings suggest that musical training may be a significant distinguishing

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factor when characterizing syllable discrimination abilities among individuals with dyslexia.
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Yet, it is important to also consider the subgroup of individuals with dyslexia, both with (n =
5) and without (n = 5) musical training, who were unable to reliably discriminate certain
syllable continua in order to complete the task(s). The majority of these individuals
exhibited difficulty with discrimination of the /ga/-/ka/ continuum which varied in temporal
information (n = 8), though this difficulty was observed in all three syllable continua. This
suggests a profound deficit in discriminating acoustic cues within syllable stimuli in
approximately 28% of the adults with dyslexia in the present study regardless of musical
training experience. This incidence (28%) is similar to previous reports of poor speech
perception skills in dyslexia (Adlard & Hazan, 1998) and difficulty discriminating temporal
cues (Overy et al., 2003). Taken together, the findings of this study support the notion that
dyslexia is characterized by individual variability with regard to the severity of deficits in
processing spectral and temporal components of speech (Law et al., 2014; Liberman, 1985),
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even among those who have had musical training.

Although enhanced speech processing abilities were predicted in musicians with dyslexia,
some weaknesses were also hypothesized for this group due to their persistent literacy
difficulties. At the group level, weaknesses were found specifically for the distinction of
amplitude envelope cues, a temporal feature critical to speech perception (Cutler, 1994;
Rosen, 1992). Considerable evidence has been put forth for temporal processing deficits in
individuals with dyslexia specific to perception of amplitude rise time and slow-rate
modulations captured by the amplitude envelope (Goswami et al., 2002; Lorenzi et al., 2000;
Rocheron et al., 2002; Talcott et al., 2000). This line of work has been further reflected by
weaknesses in both musicians and nonmusicians with dyslexia in the detection of amplitude
rise time cues within a non-linguistic context (Bishop-Liebler et al., 2014). Thus, relative to
typical musicians, the observed weaknesses in discrimination of the /bɑ/-/wɑ/ contrast
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(characterized by changes in amplitude envelope cues) in the present sample of musicians

with dyslexia as well as nonmusicians with dyslexia is in line with prior evidence. Despite
the specialized speech perception abilities that these musicians with dyslexia exhibited for
the other two syllable continua, discrimination of amplitude information within speech
syllables proved to be an area of difficulty and may be associated with their persistent
literacy difficulties.

While this study has found a distinct profile of auditory sequencing and speech processing
abilities in musicians with dyslexia that is specialized relative to individuals with dyslexia
who have not had musical training, a significant question remains: why do these individuals
still have reading difficulties? Only individuals with persistent reading difficulties were
recruited in the current study within the groups with dyslexia. Therefore, the musicians with
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dyslexia described presently are those whose reading difficulties have not been remediated
over time despite long-term musical training. These persistent deficits are also evident in the
literacy-related measures acquired within this study, as all participants with dyslexia
obtained a standardized score below 90 on at least one of the reading tests and mean
performance on all reading measures was within the low average range.

Thus, despite the specialized auditory sequencing and speech processing abilities found in
musicians with dyslexia, these individuals do not show significantly better reading abilities

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than the nonmusical controls. Musicians and nonmusicians with dyslexia did not
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significantly differ on most of the literacy-related and working memory measures; musicians
with dyslexia only achieved better scores than nonmusicians with dyslexia on the sight word
reading efficiency subtest, which measures the ability to read familiar, well-known sight
words as quickly as possible within one minute. This could point to possible compensatory
strengths within these musicians with dyslexia. Nonetheless, it remains unclear whether
these groups may differ in reading fluency or comprehension abilities, as these measures
were not included in the present study design and should be explored in future research.
Regardless, the source of persistent literacy difficulties among these adults with dyslexia
remains unclear; whether it may be deficits in auditory working memory as suggested by
Weiss and colleagues (2014) that were also found presently in both groups with dyslexia, or
perhaps the observed difficulties in discrimination of syllables characterized by changes in
amplitude envelope cues (i.e., the /bɑ/-/wɑ/ contrast). It is also possible that the perceptual
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difficulties observed in these individuals with persistent reading difficulties are due to a
general underlying difficulty with perceptual anchoring, as suggested by Ahissar and
colleagues (Ahissar et al., 2006; Oganian & Ahissar, 2012). In addition, the variance in
performance on phonological and literacy measures among those with dyslexia in this
sample supports the multidimensional view of dyslexia, which suggests a complex interplay
between the factors that are associated with reading difficulties (Pennington, 2006).
Furthermore, our findings suggest that non-literacy based experience, namely musical
training, may also contribute to reading skill development. Overall, future longitudinal
investigation is necessary to address these remaining unknowns and disentangle the extent to
which musical training may directly modulate perceptual abilities, including whether it may
serve to prevent persistent difficulties associated with dyslexia.

Taken together, this collective profile of auditory sequencing, speech processing, and literacy
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skills in musicians with dyslexia calls into question what the most significant contributing
factors may be that have shaped these processing abilities in adulthood. Two longitudinal
studies to date have demonstrated literacy improvements following musical training in 8–11
year-old children with dyslexia, who received different types of musical training in each
study (Flaugnacco et al., 2015; Habib et al., 2016). Although the evidence from the present
study supports the potential of musical training to influence literacy outcomes, it remains
unclear whether musical training directly supported the specialized auditory skills found in
these adults since the present study did not employ a longitudinal design or capture these
abilities during a developmental time period. Alternatively, since previous literature has
demonstrated that not all individuals with dyslexia exhibit weaknesses in non-linguistic
auditory processing (Hamalainen et al., 2013), it is possible that these musicians with
dyslexia never had significant weaknesses in auditory processing, even from an early age.
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Based on the present study design, this study is unable to discern the answer to this question
of whether these specialized processing abilities are a direct consequence of musical training
or rather a propensity for success with musical training.

In addition, the present study is unable to address whether the persistence of dyslexia despite
long-term musical training may point to the limitations of the benefit that musical training
may offer in the literacy domain. This is particularly important to consider given the
longitudinal evidence that literacy improvements following six weeks of computerized

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rhythm-based intervention in nine-year old children with dyslexia did not significantly differ
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from the improvements of controls who participated in phonics-based intervention or even

passive controls who did not receive direct intervention (Thomson et al., 2013). Considering
that those children were nine years old and that the average age of onset of musical training
among the musicians with dyslexia in the present sample was seven years, this additionally
calls into question whether musical training may have afforded more benefit if provided at
an earlier age, concurrent with the period of rapid development of pre-literacy and literacy
skills. In addition, the type and extent of musical training may be an important factor, as this
computerized rhythm-based intervention did not lead to the same magnitude of effect as
found in the study that implemented traditional musical training for two hours per week over
30 weeks (Flaugnacco et al., 2015). Another possibility could be that musical training
concurrent with language and literacy-based intervention may lead to maximal benefits, such
as the music-based intervention employed by Habib and colleagues that directly integrated
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shared concepts between music and language to target dyslexia-specific goals (Habib et al.,
2016). Thus, further research is needed within an earlier developmental time period to more
fully assess the benefits of administering combined music and reading-related instruction/
interventions on long-term literacy skill development and determine which approach may be
the most effective.

Alternatively, it is possible there are factors that were not addressed directly within the
present study that significantly contributed to these individuals’ successes with musical
training. These may include (but are not limited to) the following: first, participants’ familial
support and/or resources, such as remediation history or home literacy environment in
childhood, given the high socioeconomic status overall of the present sample. Second,
personal factors such as resilience or perseverance may also be significant traits among these
musicians with dyslexia that have guided them to success with music, given the unique
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strengths that have been identified in some individuals with dyslexia (Davis, 2010). Third, it
is also possible that these individuals have developed a compensatory strategy that is
advantageous for auditory processing and distinct from others with dyslexia (i.e., those who
have not had musical training). More detailed documentation of each individual’s treatment
history would have been valuable to determine the extent to which therapy experience
supported the development of compensatory mechanisms, and how these experiences may
relate to musical training status as well as non-speech and speech processing abilities. It is
also important to note that although musicians and nonmusicians with dyslexia in this study
did not differ in their average age of diagnosis, this average age was 17 years, which seems
relatively late in development to receive a diagnosis. By implication, it seems likely that
many of these individuals probably did not receive literacy-specific intervention during
literacy onset in childhood, although they may have received general classroom support.
Author Manuscript

This is conceivable given the present standard clinical diagnostic procedures in the UK
(DfES, 2005), though it raises a question of whether and how these individuals with dyslexia
managed their academics with minimal support services until college, and how age of
diagnosis may relate to long-term language, literacy, and musical abilities. Lastly, the
present sample was not large enough to evaluate whether specific attributes of the
specialized non-speech and speech-specific processing among musicians in the present
sample may be associated with the type of instrument studied. This could be of interest to

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 Zuk et al. Page 19

explore in the future, since instrument-specific effects have been proposed previously (Carey
Author Manuscript

et al., 2015). For example, string instrumentalists have demonstrated particular

specialization in the pitch domain (Koelsch et al., 1999), and percussion instrumentalists
have been characterized by specific expertise in temporal features such as timing and rhythm
(Patel, 2012). While the present study advances the extant evidence investigating auditory
processing abilities in musicians with dyslexia, these considerations are to be addressed in
future research pursuits.

In conclusion, the present study provides further specification of auditory sequencing and
speech processing abilities that characterize musicians with dyslexia who show persistent
reading difficulties. Musicians with dyslexia have demonstrated specialized processing
abilities for auditory sequencing and multiple speech-specific contexts, for measures with no
significant differences relative to typical musicians, and performed significantly more poorly
than typical musicians only for the discrimination of syllables that varied in amplitude
Author Manuscript

envelope cues. Our findings suggest that it is important to account for musical training in the
investigation of auditory processing skills in individuals with dyslexia, for musical
involvement may shape long-term auditory processing abilities. Furthermore, implications
are evident for the potential of musical training to support specific aspects of auditory
processing in individuals with dyslexia. However, more developmental and longitudinal
studies are needed to determine whether these advantages are indeed a direct result of
musical training as opposed to predispositions for success with music, whether there is a
specific profile of abilities within those with dyslexia that will benefit most from musical
training, and whether a combined music and literacy-based instruction/intervention may be
the most effective approach. Nevertheless, this work suggests that it is important to maintain
music programs within the grade school curriculum, and advocates for children with
dyslexia to continue to participate in music in addition to direct evidence-based literacy
Author Manuscript

support.

Acknowledgements
We thank all participants who took part in this study. We also thank Jennifer Minas, Michael Figuccio, and Barbara
Peysakovich for their contributions to early analyses; as well as Joseph Sanfilippo and Jacqueline Kenitz for their
editing assistance. This research was supported by the GRAMMY Foundation, the William F. Milton Funds, and
the National Institute of Health Institutional National Research Service Award (NIH T32 DC000038–22 to Zuk).

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Figure 1.
Experimental Stimuli: The spectrograms show the end points of the three continua /bɑ/-/
dɑ/, /bɑ/-/wɑ/, /ɡɑ/-/kɑ/). The onset value of the second formant in the /bɑ/-/dɑ/ continuum
varied from 800– 1600 Hz. The duration of the formant transition in the /bɑ/-/wɑ/
continuum varied from 25–97 ms. The Voice Onset Time (VOT) of the first formant in the /
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ɡɑ/-/kɑ/ continuum varied from 10–60 ms.

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Figure 2.
Performance accuracy on the tone sequence task displayed by group (open circles:
TYPMUS, filled squares: DYSMUS, filled triangle: DYSNonMUS) for the two-, three-, and
four-tone sequences. Gray lines indicate chance (2-tones, 0.25; 3-tones, 0.125; 4-tones,
0.0625).
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Figure 3.
Outcome for the three syllable continua on the syllable task. Mean Relative Threshold
Indices (RTI) by group is displayed (dark gray: TYPMUS, light gray: DYSMUS, gray:
DYSNonMUS) for each syllable continua; error bars indicate standard error. Higher RTI
scores represent better discrimination. Significance indicated by *p < 0.05 and **p < 0.01
for post-hoc Games-Howell comparisons between groups.
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Table 1

Overview of musical experience and type of instrument for TYPMUS and DYSMUS
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TYPMUS DYSMUS Sig (p-value)

Mean ± SD Mean ± SD Two-tailed
Musical Characteristics
Age began musical training 7.25 ± 2.67 7.35 ± 3.12 0.92
Years of musical training 12.88 ± 4.18 13.18 ± 2.63 0.81

Type of Musical
Instrument Number of Adults Number of Adults
Woodwinds 3 5
String 1 4
Brass 0 3
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Keyboard 4 1
Percussion 0 1
Voice 9 5
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Table 2

Attributes of the syllable continua and calculation of the relative threshold index (RTI)
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Syllable Pair Reference Syllable Original Threshold Relative Threshold Index

/bɑ/-/dɑ/
800Hz − /bɑ/ (800Hz) x Hz 1 – [x Hz - 800 / (1600Hz – 800Hz)]
1600Hz

/bɑ/-/wɑ/
/bɑ/ (25ms) x ms 1− [x ms - 25 / (97ms – 25ms)]
25ms – 97ms

/ɡɑ/-/kɑ/ 1 − [x ms - 10 / (60ms – 10ms)]

/ɡɑ/ (10ms) x ms
10ms – 60ms
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Table 3

Group characteristics as outlined by standardized measures of phonological processing, reading, and spelling
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TYPMUS DYSMUS DYSNonMUS F (max df = 2,49)

Mean ± SD Mean ± SD Mean ± SD
Group Characteristics
WRIT Verbal IQ 117.78 ± 11.78 110.11 ± 8.17 107.19 ± 12.07 a
5.08*
Nonverbal IQ 114.89 ± 7.23 113.45 ± 5.34 111.34 ± 6.54 1.19
Sight Word
TOWRE Efficiency 105.94 ± 8.44 87.42 ± 7.60 80.43 ± 6.62 b
50.23*
Phonemic
Decoding 114.35 ± 6.78 87.52 ± 6.04 84.12 ± 8.81 c
89.30***
WRAT Reading 113.88 ± 6.41 99.36 ± 7.19 95 ± 7.28 c
33.76***
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Spelling 113.47 ± 6.34 97 ± 7.13 97.18 ± 6.75 c

33.56***
Digit Memory 109.82 ± 14.24 89.47 ± 10.25 85.68 ± 11.93 c
19.21***
CTOPP Elision 107.06 ± 3.98 91.06 ± 12.98 87.18 ± 16.12 c
12.75***
Blending 115.59 ± 8.27 102.22 ± 13.97 95 ± 13.67 c
12.08***
Phonological
Awareness
Composite 113.59 ± 6.01 95.83 ± 12.78 90.125 ± 12.47 c
21.15***
Rapid Digit
Naming 110.29 ± 7.39 96.44 ± 13.82 87.81 ± 11.69 c
16.60***
Rapid Letter
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Naming 106.76 ± 11.45 87.72 ± 10.69 83.87 ± 11.84 c

19.75***
Rapid Naming
Composite 110.24 ± 10.18 90.5 ± 13.81 82.94 ± 13.58 c
20.76***

*
p < 0.05
**
p < 0.01
***
p < 0.001
a
Games-Howell post-hoc tests on one-way ANOVA by group found that TYPMUS performed significantly better than DYSNonMUS; DYSMUS
and DYSNonMUS did not significantly differ
b
Games-Howell post-hoc tests on one-way ANOVA by group found that TYPMUS performed significantly better than DYSMUS, who are in turn
better than DYSNonMUS
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c
Games-Howell post-hoc tests on one-way ANOVA by group found that TYPMUS performed significantly better than DYSMUS and
DYSNonMUS; DYSMUS and DYSNonMUS did not significantly differ

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Table 4

Discrimination thresholds for syllable continua by group as described by the Relative Threshold Index (RTI)
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TYPMUS DYSMUS DYSNonMUS F (max df = 2,49)

RTI RTI RTI

/bɑ/-/dɑ/ 0.43 0.41 0.26 b

9.20***
/bɑ/-/wɑ/ 0.68 0.47 0.39 a
7.32**
/ɡɑ/-/kɑ/ 0.37 0.31 0.14 b
5.01*

*
p < 0.05
**
p < 0.01
***
p < 0.001
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a
Games-Howell post-hoc tests on one-way ANOVA by group found that TYPMUS performed significantly better than DYSMUS and
DYSNonMUS; DYSMUS and DYSNonMUS did not significantly differ
b
Games-Howell post-hoc tests on one-way ANOVA by group found that TYPMUS did not significantly differ from DYSMUS; both TYPMUS and
DYSMUS were better than DYSNonMUS
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