Methods
2.
Children were required to read words one by one on the screen while experimenters recorded
correct and incorrect answers using a computer keyboard. Answers were coded from audio
recordings. Each correct pronouncement of the word was coded as 1, otherwise 0. A high score
indicated higher reading proficiency.
Working Memory Task
To account for these broader cognitive influences on reading development, we included
a visuo -spatial working memory task to behaviourally assess domain -general executive
function. The visuo-spatial WM maintenance task followed the protocol used by Sato et al.
(2018)75.
Participants were presented with two coloured squares (sample stimulus) for 250ms, followed
by a fixation cross for 1000ms (retention period), where they were required to remember the
previous colours in the squares. After this, two coloured squares will reappear (same colour to
previous ones or different) with a question mark as fixation point (test stimulus). During the
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test stimulus trials, participants will indicate with a computer keyboard whether the sample and
test stimuli were the same or different. Participants will press the Tick key, when they matched
and Cross key when they did not (keys m and n were covered with stickers). Participants were
instructed to respond with their dominant hand using the index and middle fingers. The test
stimulus period lasted for 2000ms or until a response was given. After the test stimulus, a
fixation cross appeared as an inter -stimulus interval (ISI) that could last between
1250ms±150ms.
There was a total of 100 experimental blocks, 50 matched blocks (row A in Fig ure 3) and 50
unmatched blocks (row B in Figure 3) and the total duration of the task was 7 minutes. At the
beginning of the task, the experimenter showed an image like Figure 2 and instructed the
participant to press the correspondent keys during test stimulus for matched and unmatched
trials. Before starting the experimental blocks, children completed a training phase where they
practiced and familiarized themselves with the proc edure. The training phase included 10
blocks of 5 matched and 5 unmatched trials.
Figure 3
Stimuli sequence for the working memory maintenance task or match-to-sample task from Sato
et al. (2018)75.
Note. A Example of matching blocks (congruent). B Example of unmatching blocks
(incongruent).
We focused on calculating accuracy and reaction times (RT) during the test stimulus phase. To
assess each participant’s performance in the task, accuracy was calculated as the percentage of
correct responses out of the total number of trials (both matched and unmatched blocks) and
RT were measured for correct trials only. Children were required to perform a minimum of
50% accuracy to be included in the analysis. An inverse efficiency score (IES) was computed
by dividing mean RT by accuracy, such that lower scores indicate more efficient performance.
This IES was used as the overall variable referred to as WM Score.
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Tapping Task
This task combines elements from Kalashnikova et al. (2021) 21, and the Expressive
Rhythm: Metronome task by Thomson and Goswami (2008)76. There were two types of trials:
Paced and Unpaced.
Paced trials. T he auditory stimuli used was a 20 -second sequence of pure tone events, each
lasting 10 milliseconds and occurring at a rate of 2Hz (500ms). Unpaced trials. Each Paced
trial was followed by a 20 -second sequence without auditory stimulus, during which
participants continued tapping at the same rate as in the Paced trials. A fixation cross appeared
on the screen during the trial. After each unpaced trial, a feedback screen displayed the
message: “Well done!.” This block of paced and unpaced trials was repeated twice. Each block
started with a countdown to prepare participants for their response (see Figure 4).
Figure 4
Visual representation of a block from the tapping task.
Participants were instructed to tap along with a metronome beat for a specific duration, aiming
to synchronise their taps as closely as possible to the metronome’s tempo. First, participants
completed a familiarization phase where the experimenter explained the task and its
procedures. The experimenter emphasized that they had to tap in sync with the sound and to
maintain the same rhythm even after the sound stopped, continuing until the cross on the screen
disappeared. The experimenter mimicked the sound and tapping as in the experimental trials,
and children were asked to tap along with their index finger on the desk. When participants
were comfortable and understood the procedure, we proceeded with the practice phase, where
they completed one block of paced and unpaced trials while the Space bar on the computer
keyboard recorded taps as input by the software. Following this, participants completed two
blocks of paced (40 s) and unpaced trials (40 s).
To ensure the analysis included only taps that were both preceded and followed by metronome
events, the first and last taps in the sequence were removed. We first calculated the mean inter-
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tap interval (ITI) for each participant in both types of trials. These scores were computed by
calculating the average time interval in milliseconds between consecutive taps for each
participant, following the metronome at a rate of 500 ms, or a tap every 500 ms. Then we
calculated the standard deviation (SD) of the ITIs for each participant in both trial types.
For the current study, we only calculated the SD of the ITIs only for paced trials which we will
refer as Tapping Variability. This measure was used as an index of inter -subject variability in
performance, capturing the consistency of tapping rates when listening to a metronome beat. A
lower Tapping Variability indicated greater consistency (i.e., more regular tapping intervals),
while a higher value indicated greater inconsistency (i.e., more variable tapping intervals).
Speech Processing Task
While EEG responses were recorded continuously, children listened to a 5 -minute
adaptation of The Gingerbread Man story (300s; 682 words) read by a male speaker and
presented through speakers. To maintain children’s engagement and minimize eye movement
artifacts, a story -related image was presented and remained on screen every 30 -40s for the
duration of the story. The images were designed to encourage to visually focus on the screen
while listening to the story. Caregivers accompanied their children if the child requested their
presence. They were instructed not to intervene or engage with the child during the task.
Envelope Extraction
Auditory envelope extraction, EEG processing and MI analyses were conducted using
MATLAB. The auditory signal was first divided into eight frequency bands, spaced equally on
a cochlear frequency map between 100 Hz and 8000 Hz, using a filter bank 77. Each band was
bandpass filtered using third-order Butterworth filters and the Hilbert transform was applied to
obtain the analytic signal. The envelope for each band was computed as the absolute value of
its analytic signal. These narrow -band envelopes w ere then averaged to produce a single
broadband envelope. The resulting envelope was bandpass filtered between 0.5 and 8 Hz to
match EEG preprocessing and then resampled to 128 Hz. Finally, the envelope was then
normalized using a Gaussian copula transform 78 and truncated to remove the 500 ms initial
offset responses79.
EEG Processing
The raw EEG data were first loaded, bad channels were rejected based on visual
inspection, data was re-referenced to average and bandpass filtered between 0.2 Hz and 40 Hz.
Independent component analysis (ICA) was applied to the filtered data (for this step, bandpass-
filtered between 1 and 40 Hz to minimise slow drifts and improve components decomposition)
to isolate and remove artefactual components related to vertical (blinks) and horizontal eye
movements and heartbeat artefacts . Components associated with electrooculography (EOG)
artifacts were detected and rejected using automated methods (based on high correlation with
the EOG), any additional components were identified and rejected based on visual inspection
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and correlation maps. ICA was then applied to the original 0.2 -40 Hz filtered data. For MI
analysis, we used the FieldTrip Toolbox80 in MATLAB. EEG data was down-sampled to 128
Hz to match the speech envelope sampling rate and bandpass filtered using third -order
Butterworth filters (0.5–8 Hz) to focus on the key frequency contributors of cortex -to-speech
envelope tacking. The EEG data w ere then segmented and aligned with the stimulus,
compensating for constant trigger delay (26 ms) between presentation of s timulus and EEG
recording times. The Hilbert transform was applied to extract the analytic signal, and the real
and imaginary components were normalized using a Gaussian-Copula transformation78.
Channel Selection
Since temporal and parietal regions are known to be involved in speech processing and
have been shown to be most sensitive to speech envelope tracking and MI analysis, we focused
the MI analysis on channels over these areas. Specific channels were chosen based on prior
research demonstrating their role in cortical speech tracking 44. For the left hemisphere these
included FT7, FC5, FC3, T7, C5, C3, TP7, CP5, CP3, P7, P5, P3, P9, and for the right
hemisphere, FT8, FC6, FC4, T8, C6, C4, TP8, CP6, CP4, P8, P6, P4, P10 (see Figure 5). This
selection ensured that the MI analysis focused on the neural response to speech rather than
visual-motor activity.
Figure 5
Visual representation of regions of interest and EEG channel selection.
MI Analysis
We employed a Mutual Information (MI) approach to assess EEG-based neural tracking
of continuous speech. MI techniques have previously been shown to effectively capture
nonlinear neural tracking of continuous speech envelopes44.
MI between continuous EEG signals and the speech envelope was estimated using the Gaussian
copula-based MI method (GCMI)78,81. Conceptually, MI is a measure that quantifies how much
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information one signal contains about another (in this case, how much information the EEG
signals contain about the speech envelope) by measuring the reduction in uncertainty about one
signal given knowledge of the other. This approach is tightly linked to that of entropy and
accounts for linear and non -linear dependencies also being robust to differences in joint and
marginal distributions.
MI was computed across a range of time lags (0 –200 ms) by shifting the EEG signal relative
to the speech envelope in 7.8 ms steps (1 sampling point at 128 Hz). Analyses were conducted
separately for different groups of channels corresponding to scalp regio ns of interest (e.g.,
left/right temporo-parietal). Within each region, MI was estimated for each participant, channel
and lag separately. To summarize these results, we extracted the maximum MI (MaxMI) for
each channel. This MaxMI value represents the highest MI observed for that channel across all
time lags, indicating the time lag at which the EEG signal aligns most strongly with the speech
envelope. Then, we selected the channel with the highest MaxMI in that region, meaning the
region-level MaxMI represents the strongest speech -brain coupling observed at any single
channel within the region . This measure highlights the peak MI response within each scalp
region, rather than an average across channels. We chose this approach because averaging
across electrodes can be biased by individual variability in head shape in young children and
electrode positioning, potentially reducing sensitivity to the strongest local speech -brain
coupling.
MI variables for further analysis included MI analyses separately for the left hemisphere and
right hemisphere, which we will refer as to Left-Hemispheric CTS (LH CTS) and Right-
Hemispheric CTS (RH CTS) respectively. Additionally, we calculated how much greater the
cortical channels in the left hemisphere track speech in comparison to the right hemisphere,
which we will refer to as Left -Lateralised CTS (subtracting left hemisphere MaxMI from the
right hemisphere MaxMI values within a participant) as we expect the frontotemporal language
network to become increasingly more left-lateralized with age9.
Analysis Plan
Our analysis plan was structured to address the study’s core hypotheses. Before we
modelled any relationships between our demographic, questionnaire, behavioural and neural
measures, we conducted group differences analyses on all key variables. Due to non -normal
distributions, we used Mann –Whitney U tests for the group comparisons and controlled for
multiple comparisons using false discovery rate (FDR) correction . These comparisons aimed
to determine whether the Music group exhibited enhanced profiles in cognitive, be havioural,
and neural measures1,8,51, consistent with our first two hypotheses.
To assess the contribution of cognitive and neural variables to reading performance, we
employed a regularised linear ridge regression model 82,83. The model included standardized
demographic (Age, Gender), questionnaire (SES, Musicality, Executive Function), behavioural
(P A, WM, Tapping Variability), neural predictors ( Left and Right-hemispheric CTS) and their
interaction with Musicality scores. Reading scores were logit -transformed after min –max
scaling to address non -normal distribution, as confirmed by visual inspection of histograms,
Q-Q plots, and skewness values. This transformation expands the scale to the full real number
line, imp roving linearity and normality assumptions for regression. Ridge regression was
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chosen over ordinary least squares due to moderate to high correlations among predictors (see
Supplementary Information - Figure 1 and Figure 2 ), which raised concerns about
multicollinearity. By applying L2 regularization, ridge regression penalizes large coefficients,
reducing variance and improving model stability, thereby preventing overfitting, and enhancing
generalizability. Model assumptio ns were thoroughly evaluated: the Durbin -Watson statistic
indicated no significant autocorrelation of residuals; the Breusch -Pagan test indicated no
heteroskedasticity; residuals approximated normality based on Q -Q plots; Variance Inflation
Factors were within acceptable ranges for most predictors; and no extreme outliers were
observed, supporting the use of ridge regression.
The aim was to determine whether language-specific mechanisms (PA and CTS) could explain
the influence of musical engagement on reading ability, consistent with prior literature 13,41,43.
To test whether PA and CTS mediate the effect of musical training on reading, we performed
parallel mediation analyses using the Pingouin statistical package in Python 84. This approach
allowed us to estimate indirect effects using non-parametric bootstrapping (20,000 resamples),
which does not assume the normality of the sampling distribution of indirect paths. The
mediation model included Musicality scores as the independent variable, Reading accuracy as
the dependent variable, PA, Left and Right-hemispheric CTS as parallel mediators, and Age,
Gender, SES, Executive Function and WM scores as covariates.
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References
1 Moreno, S., Marques, C., Santos, A., Santos, M., Castro, S. L., & Besson, M. (2009). Musical
training influences linguistic abilities in 8 -year-old children: more evidence for brain
plasticity. Cerebral cortex (New York, N.Y . : 1991), 19(3), 712 –723.
https://doi.org/10.1093/cercor/bhn120
2 Dos Santos-Luiz, C., Mónico, L. S. M., Almeida, L. S., & Coimbra, D. (2016). Exploring the
long-term associations between adolescents’ music training and academic achievement.
Musicae Scientiae, 20(4), 512–527. https://doi.org/10.1177/1029864915623613
3 Janurik, M., & Józsa, K. (2022). Long-Term Impacts of Early Musical Abilities on Academic
Achievement: A Longitudinal Study. Journal of Intelligence, 10(3), 36.
https://doi.org/10.3390/jintelligence10030036
4 Dehaene, S. (2009). Reading in the brain: The new science of how we read. Penguin.
5 Ivanova, M. V ., Zhong, A., Turken, A., Baldo, J. V ., & Dronkers, N. F. (2021). Functional
Contributions of the Arcuate Fasciculus to Language Processing. Frontiers in human
neuroscience, 15, 672665. https://doi.org/10.3389/fnhum.2021.672665
6 Pleisch, G., Karipidis, I. I., Brauchli, C., Röthlisberger, M., Hofstetter, C., Stämpfli, P.,
Walitza, S., & Brem, S. (2019). Emerging neural specialization of the ventral
occipitotemporal cortex to characters through phonological association learning in
preschool children. NeuroImage, 189, 813–831.
https://doi.org/10.1016/j.neuroimage.2019.01.046
7 Olulade, O. A., Seydell-Greenwald, A., Chambers, C. E., Turkeltaub, P. E., Dromerick, A. W.,
Berl, M. M., Gaillard, W. D., & Newport, E. L. (2020). The neural basis of language
development: Changes in lateralization over age. Proceedings of the National Academy
of Sciences of the United States of America, 117(38), 23477 –23483.
https://doi.org/10.1073/pnas.1905590117
8 Zatorre, R. J., Belin, P., & Penhune, V . B. (2002). Structure and function of auditory cortex:
music and speech. Trends in cognitive sciences, 6(1), 37 –46.
https://doi.org/10.1016/s1364-6613(00)01816-7
9 Rosselli, M., Ardila, A., Matute, E., & Vélez-Uribe, I. (2014). Language Development across
the Life Span: A Neuropsychological/Neuroimaging Perspective. Neuroscience journal,
2014, 585237. https://doi.org/10.1155/2014/585237
10 Nenert, R., Allendorfer, J. B., Martin, A. M., Banks, C., Vannest, J., Holland, S. K., &
Szaflarski, J. P. (2017). Age -related language lateralization assessed by fMRI: The
effects of sex and handedness. Brain research, 1674, 20 –35.
https://doi.org/10.1016/j.brainres.2017.08.021
11 Fedorenko, E., Behr, M. K., & Kanwisher, N. (2011). Functional specificity for high -level
linguistic processing in the human brain. Proceedings of the National Academy of
Sciences of the United States of America, 108(39), 16428 –16433.
https://doi.org/10.1073/pnas.1112937108
preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.
The copyright holder for thisthis version posted September 6, 2025. ; https://doi.org/10.1101/2025.09.05.674218doi: bioRxiv preprint
12 Kraus, N., Hornickel, J., Strait, D. L., Slater, J., & Thompson, E. (2014). Engagement in
community music classes sparks neuroplasticity and language development in children
from disadvantaged backgrounds. Frontiers in psychology, 5, 1403.
https://doi.org/10.3389/fpsyg.2014.01403
13 Flaugnacco, E., Lopez, L., Terribili, C., Montico, M., Zoia, S., & Schön, D. (2015). Music
training increases phonological awareness and reading skills in developmental
dyslexia: A randomized control trial. PLOS ONE, 10(9), e0138715.
https://doi.org/10.1371/journal.pone.0138715
14 Gordon, R. L., Fehd, H. M., & McCandliss, B. D. (2015). Does Music Training Enhance
Literacy Skills? A Meta -Analysis. Frontiers in psychology, 6, 1777.
https://doi.org/10.3389/fpsyg.2015.01777
15 Sofologi, M., Papatzikis, E., Kougioumusical trainingzis, G., Kosmidou, E., Klitsioti, A.,
Droutme, A., Sourbi, A. A., Chrisostomou, D., & Efstratopoulou,
M. (2022). Effectiveness of Musical training on Reading Comprehension in
Elementary School Children. Is There an Associative Cognitive Benefit? Frontiers in
Education, 7, Article 875511. https://doi.org/10.3389/feduc.2022.875511
16 Patel, A. D. (2011). Why would musical training benefit the neural encoding of speech? The
OPERA hypothesis. Frontiers in Psychology, 2, Article 142.
https://doi.org/10.3389/fpsyg.2011.00142
17 Fiveash, A., Bedoin, N., Gordon, R. L., & Tillmann, B. (2021). Processing rhythm in speech
and music: Shared mechanisms and implications for developmental speech and
language disorders. Neuropsychology, 35(8), 771 –791.
https://doi.org/10.1037/neu0000766
18 Patscheke, H., Degé, F., & Schwarzer, G. (2016). The Effects of Training in Music and
Phonological Skills on Phonological Awareness in 4 - to 6 -Year-Old Children of
Immigrant Families. Frontiers in psychology, 7, 1647.
https://doi.org/10.3389/fpsyg.2016.01647
19 Ziegler, J. C., & Goswami, U. (2005). Reading acquisition, developmental dyslexia, and
skilled reading across languages: a psycholinguistic grain size theory. Psychological
bulletin, 131(1), 3–29. https://doi.org/10.1037/0033-2909.131.1.3
20 Woodruff Carr, K., White-Schwoch, T., Tierney, A. T., Strait, D. L., & Kraus, N. (2014). Beat
synchronization predicts neural speech encoding and reading readiness in preschoolers.
Proceedings of the National Academy of Sciences of the United States of Ame rica,
111(40), 14559–14564. https://doi.org/10.1073/pnas.1406219111
21 Kalashnikova, M., Burnham, D., and Goswami, U. (2021). Rhythm discrimination and
metronome tapping in 4-year-old children at risk for developmental dyslexia. Cog. Dev.
60:101129. https://doi.org/10.1016/j.cogdev.2021.101129
22 Zendel, B. R., West, G. L., Belleville, S., & Peretz, I. (2019). Musical training improves the
ability to understand speech-in-noise in older adults. Neurobiology of aging, 81, 102 –
115. https://doi.org/10.1016/j.neurobiolaging.2019.05.015
preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.
The copyright holder for thisthis version posted September 6, 2025. ; https://doi.org/10.1101/2025.09.05.674218doi: bioRxiv preprint
23 Tierney, A. T., Krizman, J., & Kraus, N. (2015). Music training alters the course of adolescent
auditory development. Proceedings of the National Academy of Sciences of the United
States of America, 112(32), 10062–10067. https://doi.org/10.1073/pnas.1505114112
24 Li, Q., Wang, X., Wang, S., Xie, Y ., Xie, Y ., & Li, S. (2020). More Flexible Integration of
Functional Systems After Musical training in Young Adults. IEEE transactions on
neural systems and rehabilitation engineering: a publication of the IEEE Engineering
in Medicine and Biology Society, 28(4), 817 –824.
https://doi.org/10.1109/TNSRE.2020.2977250
25 Schellenberg, E. G., & Lima, C. F. (2024). Music Training and Nonmusical Abilities. Annual
review of psychology, 75, 87 –128. https://doi.org/10.1146/annurev -psych-032323-
051354
26 Bigand, E., & Tillmann, B. (2022). Near and far transfer: Is music special?. Memory &
Cognition, 50(2), 339-347. https://doi.org/10.3758/s13421-021-01226-6
27 Giraud, A. L., & Poeppel, D. (2012). Cortical oscillations and speech processing: emerging
computational principles and operations. Nature neuroscience, 15(4), 511 –517.
https://doi.org/10.1038/nn.3063
28 Obleser, J., & Kayser, C. (2019). Neural Entrainment and Attentional Selection in the
Listening Brain. Trends in cognitive sciences, 23(11), 913 –926.
https://doi.org/10.1016/j.tics.2019.08.004
29 Issa, M. F., Khan, I., Ruzzoli, M., Molinaro, N., & Lizarazu, M. (2024). On the speech
envelope in the cortical tracking of speech. NeuroImage, 297, 120675.
https://doi.org/10.1016/j.neuroimage.2024.120675
30 Ghitza O. (2011). Linking speech perception and neurophysiology: speech decoding guided
by cascaded oscillators locked to the input rhythm. Frontiers in psychology, 2, 130.
https://doi.org/10.3389/fpsyg.2011.00130
31 Atanasova, T., Gross, J., Keitel, A., & Rimmele, J. M. (2025). The involvement of
endogenous brain rhythms in speech processing. PsyArXiv Preprints.
https://doi:10.31234/osf.io/rukwp_v1
32 Lalor, E., & Nidiffer, A. (2025, May 20). On the generative mechanisms underlying the
cortical tracking of natural speech: a position paper.
https://doi.org/10.31219/osf.io/xf8ay_v1
33 Whiteford, K.L., Baltzell, L.S., Chiu, M. et al. Large -scale multi -site study shows no
association between musical training and early auditory neural sound encoding. Nat
Commun 16, 7152 (2025). https://doi.org/10.1038/s41467-025-62155-5
34 Goswami, U. (2011). A temporal sampling framework for developmental dyslexia. Trends in
Cognitive Sciences, 15(1), 3–10. https://doi.org/10.1016/j.tics.2010.10.001
35 Leong, V ., Stone, M. A., Turner, R. E., & Goswami, U. (2014). A role for amplitude
modulation phase relationships in speech rhythm perception. The Journal of the
Acoustical Society of America, 136(1), 366–381. https://doi.org/10.1121/1.4883366
preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.
The copyright holder for thisthis version posted September 6, 2025. ; https://doi.org/10.1101/2025.09.05.674218doi: bioRxiv preprint
36 Destoky, F., Bertels, J., Niesen, M., Wens, V ., Vander Ghinst, M., Rovai, A., Trotta, N.,
Lallier, M., De Tiège, X., & Bourguignon, M. (2022). The role of reading experience
in atypical cortical tracking of speech and speech -in-noise in dyslexia. NeuroImage,
253, 119061. https://doi.org/10.1016/j.neuroimage.2022.119061
37 Nora, A., Rinkinen, O., Renvall, H., Service, E., Arkkila, E., Smolander, S., Laasonen, M.,
& Salmelin, R. (2024). Impaired Cortical Tracking of Speech in Children with
Developmental Language Disorder. The Journal of neuroscience : the official journal
of the Society for Neuroscience, 44(22), e2048232024.
https://doi.org/10.1523/JNEUROSCI.2048-23.2024
38 Goswami, U. (2018). A neural basis for phonological awareness? An oscillatory temporal -
sampling perspective. Current Directions in Psychological Science, 27(1), 56 –63.
https://doi.org/10.1177/0963721417727520
39 Tierney, A., & Kraus, N. (2013). Music training for the development of reading
skills. Progress in brain research, 207, 209–241. https://doi.org/10.1016/B978-0-444-
63327-9.00008-4
40 Rogachëv, A., & Sysoeva, O. (2024). Neural tracking of natural speech in children in relation
to their receptive speech abilities. Cognitive Systems Research, 86, Article 101236.
https://doi.org/10.1016/j.cogsys.2024.101236
41 Sousa, J., Martins, M., Torres, N., Castro, S. L., & Silva, S. (2022). Rhythm but not melody
processing helps reading via phonological awareness and phonological memory.
Scientific reports, 12(1), 13224. https://doi.org/10.1038/s41598-022-15596-7
42Abrams, D. A., Nicol, T., Zecker, S., & Kraus, N. (2009). Abnormal cortical processing of
the syllable rate of speech in poor readers. The Journal of neuroscience: the official
journal of the Society for Neuroscience, 29(24), 7686 –7693.
https://doi.org/10.1523/JNEUROSCI.5242-08.2009
43 Musacchia, G., Sams, M., Skoe , E., & Kraus, N. (2007). Musicians have enhanced
subcortical auditory and audiovisual processing of speech and music. Proceedings of
the National Academy of Sciences of the United States of America, 104(40), 15894 –
15898. https://doi.org/10.1073/pnas.0701498104
44 De Clercq, P., Vanthornhout, J., Vandermosten, M., & Francart, T. (2023). Beyond linear
neural envelope tracking: a mutual information approach. Journal of neural
engineering, 20(2), 10.1088/1741 -2552/acbe1d. https://doi.org/10.1088/1741-
2552/acbe1d
45 Janurik, M., Surján, N., & Józsa, K. (2022). The Relationship between Early Word Reading,
Phonological Awareness, Early Music Reading and Musical Aptitude. Journal of
Intelligence, 10(3), 50. https://doi.org/10.3390/jintelligence10030050
46 Politimou, N., Dalla Bella, S., Farrugia, N., & Franco, F. (2019). Born to Speak and Sing:
Musical Predictors of Language Development in Pre -schoolers. Frontiers in
psychology, 10, 948. https://doi.org/10.3389/fpsyg.2019.00948
47 Turesky, T. K., Escalante, E., Loh, M., & Gaab, N. (2025). Longitudinal trajectories of brain
development from infancy to school age and their relationship to literacy development.
preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.
The copyright holder for thisthis version posted September 6, 2025. ; https://doi.org/10.1101/2025.09.05.674218doi: bioRxiv preprint
bioRxiv : the preprint server for biology, 2024.06.29.601366.
https://doi.org/10.1101/2024.06.29.601366
48 Tan, F. M., Yu, J., & Goodwill, A. M. (2025). Sports participation & childhood
neurocognitive development. Developmental cognitive neuroscience, 71, 101492.
https://doi.org/10.1016/j.dcn.2024.101492
49 Sun, S., & Chen, C. (2024). The Effect of Sports Game Intervention on Children's
Fundamental Motor Skills: A Systematic Review and Meta-Analysis. Children (Basel,
Switzerland), 11(2), 254. https://doi.org/10.3390/children11020254
50 Frischen, U., Schwarzer, G., & Degé, F. (2021). Music lessons enhance executive functions
in 6 -to 7 -year-old children. Learning and Instruction, 74, 101442.
https://doi.org/10.1016/j.learninstruc.2021.101442
51 Degé, F., & Schwarzer, G. (2011). The effect of a music program on phonological awareness
in preschoolers. Frontiers in Psychology, 2, Article 124.
https://doi.org/10.3389/fpsyg.2011.00124
52 Colling, L. J., Noble, H. L., & Goswami, U. (2017). Neural Entrainment and Sensorimotor
Synchronization to the Beat in Children with Developmental Dyslexia: An EEG Study.
Frontiers in neuroscience, 11, 360. https://doi.org/10.3389/fnins.2017.00360
53 Habib, M., Lardy, C., Desiles, T., Commeiras, C., Chobert, J., & Besson, M. (2016). Music
and Dyslexia: A New Musical Training Method to Improve Reading and Related
Disorders. Frontiers in psychology, 7, 26. https://doi.org/10.3389/fpsyg.2016.00026
54 Etard, O., & Reichenbach, T. (2019). Neural Speech Tracking in the Theta and in the Delta
Frequency Band Differentially Encode Clarity and Comprehension of Speech in Noise.
The Journal of neuroscience: the official journal of the Society for Neuroscience,
39(29), 5750–5759. https://doi.org/10.1523/JNEUROSCI.1828-18.2019
55 Pérez-Navarro, J., Klimovich-Gray, A., Lizarazu, M., Piazza, G., Molinaro, N., & Lallier, M.
(2024). Early language experience modulates the tradeoff between acoustic -temporal
and lexico -semantic cortical tracking of speech. iScience, 27(7), 110247.
https://doi.org/10.1016/j.isci.2024.110247
56 Lizarazu, M., Carreiras, M., Bourguignon, M., Zarraga, A., & Molinaro, N. (2021). Language
Proficiency Entails Tuning Cortical Activity to Second Language Speech. Cerebral
cortex (New York, N.Y . : 1991), 31(8), 3820 –3831.
https://doi.org/10.1093/cercor/bhab051
57Molinaro, N., & Lizarazu, M. (2018). Delta (but not theta)-band cortical entrainment involves
speech-specific processing. The European journal of neuroscience, 48(7), 2642 –2650.
https://doi.org/10.1111/ejn.13811
58 Melby-Lervåg, M., Lyster, S.-A. H., & Hulme, C. (2012). Phonological skills and their role
in learning to read: A meta -analytic review. Psychological Bulletin, 138(2), 322 –352.
https://doi.org/10.1037/a0026744
preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.
The copyright holder for thisthis version posted September 6, 2025. ; https://doi.org/10.1101/2025.09.05.674218doi: bioRxiv preprint
59 Diamanti, V ., Grande, G., Protopapas, A., Melby-Lervåg, M., & Lervåg, A. (2024). Preschool
morphological awareness and developmental change in early reading ability. Scientific
Studies of Reading, 28(6), 591–613. https://doi.org/10.1080/10888438.2024.2370843
60 Bhide, A., Power, A., & Goswami, U. (2013). A rhythmic musical intervention for poor
readers: A comparison of efficacy with a letter‐based intervention. Mind, Brain, and
Education, 7(2), 113–123. https://doi.org/10.1111/mbe.12016
61 Rautenberg, I. (2015). The effects of musical training on the decoding skills of German‐
speaking primary school children. Journal of Research in Reading, 38(1), 1 –17.
https://doi.org/10.1111/jrir.12010
62 Molinaro, N., Lizarazu, M., Lallier, M., Bourguignon, M., & Carreiras, M. (2016). Out -of-
synchrony speech entrainment in developmental dyslexia. Human brain mapping,
37(8), 2767–2783. https://doi.org/10.1002/hbm.23206
63 Rezlescu, C., Danaila, I., Miron, A., & Amariei, C. (2020). More time for science: Using
Testable to create and share behavioral experiments faster, recruit better participants,
and engage students in hands-on research. Progress in Brain Research, 253, 243-262.
64 Peirce, J., Gray, J. R., Simpson, S., MacAskill, M., Höchenberger, R., Sogo, H., Kastman,
E., & Lindeløv, J. K. (2019). PsychoPy2: Experiments in behavior made easy. Behavior
research methods, 51(1), 195–203. https://doi.org/10.3758/s13428-018-01193-y
65 Gramfort, A., Luessi, M., Larson, E., Engemann, D. A., Strohmeier, D., Brodbeck, C., Goj,
R., Jas, M., Brooks, T., Parkkonen, L., & Hämäläinen, M. (2013). MEG and EEG data
analysis with MNE -Python. Frontiers in neuroscience, 7, 267.
https://doi.org/10.3389/fnins.2013.00267
66 Politimou, N., Stewart, L., Müllensiefen, D., & Franco, F. (2018). Music@Home: A novel
instrument to assess the home musical environment in the early years. PloS one, 13(4),
e0193819. https://doi.org/10.1371/journal.pone.0193819
67 Dolean, D., Melby -Lervåg, M., Tincas, I., Damsa, C., & Lervåg, A. (2019). Achievement
gap: Socioeconomic status affects reading development beyond language and cognition
in children facing poverty. Learning and Instruction, 63, Article 101218.
https://doi.org/10.1016/j.learninstruc.2019.101218
68 Rodriguez-Gomez, D. A., & Talero -Gutiérrez, C. (2022). Effects of music training in
executive function performance in children: A systematic review. Frontiers in
psychology, 13, 968144. https://doi.org/10.3389/fpsyg.2022.968144
69 Meixner, J.M.; Laubrock, J. Executive functioning predicts development of reading skill and
perceptual span seven years later. J. Mem. Lang. 2024, 136, 104511.
https://doi.org/10.1016/j.jml.2024.104511
70 Escobar, J. P., Espinoza, V ., & Balboa, S. (2024). Relations Between Executive Functions
and Reading Comprehension: A Study of Fourth -Grade Students with and Without
Reading Comprehension Difficulties. Brain sciences, 14(12), 1174.
https://doi.org/10.3390/brainsci14121174
preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.
The copyright holder for thisthis version posted September 6, 2025. ; https://doi.org/10.1101/2025.09.05.674218doi: bioRxiv preprint
71 Gioia, G. A., Isquith, P. K., Guy, S. C., & Kenworthy, L. (2015). BRIEF2: Behavior Rating
Inventory of Executive Function (2nd ed.). Psychological Assessment Resources.
https://doi.org/10.1037/t79467-000
72 University of Oregon (2019). 8th Edition of Dynamic Indicators of Basic Early Literacy Skills
(DIBELS®): Technical Manual. Eugene, OR: Author. Available:
https://dibels.uoregon.edu/
73 Zumeta, R. O., Compton, D. L., & Fuchs, L. S. (2012). Using Word Identification Fluency
to Monitor First -Grade Reading Development. Exceptional children, 78(2), 201 –220.
https://doi.org/10.1177/001440291207800204
74 Dolch, E. W. (1936). A basic sight vocabulary. The Elementary School Journal, 36, 456–60.
https://doi.org/10.1086/457353
75 Sato, J., Mossad, S. I., Wong, S. M., Hunt, B. A. E., Dunkley, B. T., Smith, M. L., Urbain,
C., & Taylor, M. J. (2018). Alpha keeps it together: Alpha oscillatory synchrony
underlies working memory maintenance in young children. Developmental cognitive
neuroscience, 34, 114–123. https://doi.org/10.1016/j.dcn.2018.09.001
76Thomson, J. M., & Goswami, U. (2008). Rhythmic processing in children with
developmental dyslexia: auditory and motor rhythms link to reading and spelling.
Journal of physiology, Paris, 102(1 -3), 120 –129.
https://doi.org/10.1016/j.jphysparis.2008.03.007
77 Smith, Z. M., Delgutte, B., & Oxenham, A. J. (2002). Chimaeric sounds reveal dichotomies
in auditory perception. Nature, 416(6876), 87–90. https://doi.org/10.1038/416087a
78 Ince, R. A., Giordano, B. L., Kayser, C., Rousselet, G. A., Gross, J., & Schyns, P. G. (2017).
A statistical framework for neuroimaging data analysis based on mutual information
estimated via a gaussian copula. Human brain mapping, 38(3), 1541 –1573.
https://doi.org/10.1002/hbm.23471
79 Park, H., Ince, R. A. A., Schyns, P. G., Thut, G., & Gross, J. (2018). Representational
interactions during audiovisual speech entrainment: Redundancy in left posterior
superior temporal gyrus and synergy in left motor cortex. PLoS biology, 16(8),
e2006558. https://doi.org/10.1371/journal.pbio.2006558
80 Oostenveld, R., Fries, P., Maris, E., & Schoffelen, J. M. (2011). FieldTrip: Open source
software for advanced analysis of MEG, EEG, and invasive electrophysiological data.
Computational intelligence and neuroscience, 2011, 156869.
https://doi.org/10.1155/2011/156869
81 Keitel A, Gross J, Kayser C (2018) Perceptually relevant speech tracking in auditory and
motor cortex reflects distinct linguistic features. PLOS Biology 16(3): e2004473.
https://doi.org/10.1371/journal.pbio.2004473
82 Hoerl, A. E., & Kennard, R. W. (1970). Ridge Regression: Biased Estimation for
Nonorthogonal Problems. Technometrics, 12(1), 55 –67.
https://doi.org/10.1080/00401706.1970.10488634
preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.
The copyright holder for thisthis version posted September 6, 2025. ; https://doi.org/10.1101/2025.09.05.674218doi: bioRxiv preprint
83 Friedman, J., Hastie, T., & Tibshirani, R. (2010). Regularization Paths for Generalized Linear
Models via Coordinate Descent. Journal of statistical software, 33(1), 1–22.
84 Vallat, R. (2018). Pingouin: Statistics in Python. Journal of Open Source Software, 3(31),
1026. https://doi.org/10.21105/joss.01026
preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.
The copyright holder for thisthis version posted September 6, 2025. ; https://doi.org/10.1101/2025.09.05.674218doi: bioRxiv preprint