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OPEN Operatic voices engage the default

mode network in professional
opera singers
Adél Bihari 1,2*, Ádám Nárai 1,3, Boris Kleber 4, Judit Zsuga 5, Petra Hermann 1,6 &
Zoltán Vidnyánszky 1,6*

Extensive research with musicians has shown that instrumental musical training can have a profound
impact on how acoustic features are processed in the brain. However, less is known about the
influence of singing training on neural activity during voice perception, particularly in response to
salient acoustic features, such as the vocal vibrato in operatic singing. To address this gap, the present
study employed functional magnetic resonance imaging (fMRI) to measure brain responses in trained
opera singers and musically untrained controls listening to recordings of opera singers performing
in two distinct styles: a full operatic voice with vibrato, and a straight voice without vibrato. Results
indicated that for opera singers, perception of operatic voice led to differential fMRI activations in
bilateral auditory cortical regions and the default mode network. In contrast, musically untrained
controls exhibited differences only in bilateral auditory cortex. These results suggest that operatic
singing training triggers experience-dependent neural changes in the brain that activate self-
referential networks, possibly through embodiment of acoustic features associated with one’s own
singing style.

Keywords Operatic voice, Auditory processing, Musical expertise, Default mode network, fMRI

Becoming a professional musician demands extensive and persistent dedication to practice and learning over an
extended period of time, often starting in early ­childhood1,2. This pursuit entails not only developing sensorimo-
tor proficiency but also encompasses a profound transformation of sensory perceptual modalities required for
performance monitoring and motor ­adjustments3–5. Simultaneously, it engages cognitive and affective processes,
highlighting its multifaceted n­ ature6,7. This comprehensive training has been associated with several anatomi-
cally plausible adaptations in brain regions involved in sensory processing, sensorimotor integration, emotion,
and higher-level cognitive functions, causing a global impact on the structural and functional organization
­ rain8–11. Robust changes in large-scale structural and functional networks across the brain, as recently
of the b
­demonstrated12, substantiate and extend these observations, indicating that musical expertise shapes the con-
nectivity across sensory, motor, and cognitive regions, potentially enhancing the information transfer among
these neural systems even in task-free ­conditions13. These results highlight why musical training has emerged
as an excellent framework for investigating the brain’s ability to adapt and reorganize in response to experiences
and environmental d ­ emands14, with broader implications for understanding brain development, learning, and
adaptation that extend beyond the realm of music ­itself15.
While we continue to expand our understanding of how musical training may alter the brain, we remain
relatively naïve about the effects of training an instrument that’s so ubiquitous, we often take it for granted: our
voice. Drawing inferences regarding the potential effect of training one’s voice carries significant implications,
since much of the research into training-related effects has focused on how long-term exposure to specific acous-
tic stimuli can alter auditory perception, considering that observable differences are most pronounced when
the auditory input is both behaviourally relevant and actively t­ rained8,16–18. This is consistent with the "action-
perception" hypothesis, suggesting that perception is not a passive process but is actively shaped by our past

1
Brain Imaging Centre, HUN-REN Research Centre for Natural Sciences, Budapest, Hungary. 2Department of
Cognitive Science, Faculty of Natural Sciences, Budapest University of Technology and Economics, Budapest,
Hungary. 3Doctoral School of Biology and Sportbiology, Institute of Biology, Faculty of Sciences, University of
Pécs, Pécs, Hungary. 4Center for Music in the Brain, Department of Clinical Medicine, Aarhus University and The
Royal Academy of Music, Aarhus/Aalborg, Denmark. 5Department of Psychiatry, Faculty of Medicine, University
of Debrecen, Debrecen, Hungary. 6These authors jointly supervised this work: Petra Hermann and Zoltán
Vidnyánszky. *email: [email protected]; [email protected]

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experiences and a­ ctions19. Further supporting this perspective, the Theory of Event Coding (TEC) and Prinz’s
model on a common coding approach postulate that sensory inputs and motor outputs are not separately pro-
cessed in the brain but are encoded as "events" in a common representational medium, enhancing the efficiency
of interaction with the e­ nvironment20,21. The Predictive Coding of Music (PCM) m ­ odel22,23 further complements
this view by framing music perception, action, emotion, and learning as recursive Bayesian processes, where
the brain continuously minimizes prediction errors by comparing sensory input with top-down predictions.
This model illustrates how musical training, such as that received by opera singers, refines these predictive
mechanisms, leading to improved auditory and motor integration and corresponding cognitive skills through
long-term experience-dependent plasticity. A compelling demonstration of the intimate relationship between
sound production and perception is that cortical responses to musical stimuli are uniquely modulated by the
specific timbre of one’s own trained ­instrument24–28. Moreover, it has been shown that the primary laryngeal
motor cortex involves separable motor and auditory representations of vocal pitch, with discrete neural popu-
lations handling specific pitch-related tasks during speech or s­ inging29. When it comes to classical or operatic
singing, vibrato is a particularly salient feature that provides a unique opportunity to explore how singers’ net
sensorimotor experience with this singing style may shape auditory perception and the ensuing brain responses.
In Western-style operatic singing, vibrato is an essential aspect of classical vocal training that must be
­mastered30. Characterized by a quasi-sinusoidal, low-frequency modulation of the fundamental frequency or
pitch, vibrato affects all spectral timbral ­components31. Typically, it is quantified in terms of the rate and extent
of this modulation, whereas the pitch we perceive corresponds to the mean frequency of this m ­ odulation32.
It serves a crucial artistic function: it infuses the voice with richness and warmth, enhancing both resonance
and emotional e­ xpression33–35. Given its importance in classical singing training, its psychoacoustic properties,
and its artistic impact, vibrato presents an intriguing lens through which to investigate the neural mechanisms
underlying experience-dependent action-perception processes.
Our present understanding of the brain function and structural differences associated with action-perception
processes in singing training largely stems from studies that have utilized pitch matching and singing production
paradigms to investigate sensorimotor integration. These studies collectively suggest an experience-dependent
role of embodied motor and acoustic feedback control mechanisms, strongly engaging the auditory, somatosen-
sory, and inferior parietal cortices, as well as frontal brain regions associated with performance monitoring and
the ­insula36–41. With regards to the sensory modalities involved, the brain regions associated with embodied
motor control not only support tactile information transmission relevant to singing but also have important
links with corresponding auditory p ­ rocesses10,35,42–44. However, despite a steadily increasing number of studies
investigating the neural mechanisms underlying human s­ peech45, understanding how singing training might
influence perceptual processes remains an open q ­ uestion46,47.
To address this gap, the current study aims to elucidate the specific neural mechanisms by which experi-
ence and training in operatic singing interact with the neural substrates underlying its perception, leveraging
the presence or absence of vibrato as a defining element. Blood-oxygen-level-dependent (BOLD) responses
were measured to ecologically valid recordings performed by trained opera singers, highlighting two contrast-
ing vocal styles: a full operatic voice with vibrato, and a straighter voice without vibrato, which we refer to as
the "natural" style. These responses were investigated in trained opera singers and subsequently compared to
musically untrained controls. We hypothesize that listening to operatic versus natural singing styles will engage
brain regions linked to auditory perception, sensorimotor integration, and higher-level cognitive functions more
robustly in opera singers. By including musically untrained controls, we aim to determine whether comprehen-
sive operatic singing training will result in heightened neural activity in these brain regions among singers relative
to controls, following our hypothesis that specialized singing expertise has a profound impact on singing-related
auditory processing in the brain.

Methods
Participants
In this study, 32 professionally trained classical opera singers and 35 controls without prior musical or vocal
training were enrolled. None of the participants had a reported history of neurological or psychiatric disease.
Due to excessive head motion and anatomical abnormalities, two controls were excluded from further analysis,
resulting in a final sample size of N = 32 opera singers (age range: 20–52 years; age mean ± standard deviation
(SD): 39.0 ± 7.3 years; 18 female, 5 left-handed) and N = 33 controls (age range: 20–49 years; age mean ± SD:
29.0 ± 8.2 years; 22 female, 2 left-handed). Professional singers’ average age at the beginning of formal private
singing training was 17.9 years (SD = 3.9; range = 13–29 years), and their average professional singing experience
was 20.8 years (SD = 6.7, range = 6–31 years), as assessed by the translated form of the Montreal Music History
Questionnaire (MMHQ)48. All participants provided written, informed consent before participation and were
financially compensated for their time. The research protocol was designed and conducted in accordance with
the Hungarian regulations and laws, and with the Declaration of Helsinki, and was approved by the National
Institute of Pharmacy and Nutrition, Hungary (file number: OGYÉI/70,184/2017).
Data acquisition initially focused on the opera group. After the analyses revealed significant effects related
to stimuli style (natural vs. operatic), we proceeded to recruit control participants to determine if these effects
were specific to opera singers. The analysis pipeline described takes the sequential nature of data acquisition
into account.

Stimuli generation
Following the methodology described by Lévêque et al.49,50, we initially generated a diverse pool of 120 pseudor-
andomized 5-tone isochronous melodies for use across multiple studies. These melodies were derived from all 7

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modal scales and were designed with minimal tone repetition (maximum one repetition per melody). The melo-
dies were transposed into three distinct vocal ranges to accommodate four different voice types: G2–C♯3 for men
only (male-low), B3–E♯4 for both sexes (male-high and female-low), and C5–F♯5 for women only (female-high).
Melodies were recorded by two professionally trained and actively performing opera singers, one male (bari-
tone) and one female (mezzo-soprano). Each singer performed the melodies using the vowel "o" in two distinct
singing styles: "naturally" (using straight tones without vibrato) and "operatically" (using vibrato). The record-
ings were captured at a sampling rate of 48 kHz in a sound-proof recording studio. A metronome set at 80 beats
per minute and the melody played on a piano delivered through headphones aided the singers in maintaining
consistent tempo and pitch. Each recorded stimulus had an average ± SD duration of 4.03 ± 0.12 s.

Stimulus selection and preparation

For this specific experiment, we refined our stimulus selection to 48 melodies, each recorded with all stimulus
type combinations (2 singing styles × 4 voice types), resulting in a total of 384 unique stimuli. During the selec-
tion process, we focused on minimizing length differences (maximum difference of 353 ms) for each melody
across all voice types and singing styles. To ensure uniformity across conditions, melodies were trimmed at both
ends to match the length of the shortest one. Onset and offset ramps were applied using a half-Hanning window,
corresponding in length to the amount trimmed. Finally, each melody was normalized to ensure a consistent
root mean square (RMS) value across all stimuli.
To optimize the use of our specialized participant pool, the curated set of 48 melodies was designed to allow
for the investigation of multiple research questions for different studies. This approach aimed to enhance the
efficiency of data collection to explore various dimensions of vocal perception, such as the impact of singing
styles, vocal ranges, and the influence of the participants’ and singers’ sex on participants’ auditory processing.
For the current analysis, however, we focused specifically on the effect of singing style (operatic vs. natural) within
the vocal range of B3–E♯4, reproducible by both male and female participants. This range was selected to avoid
potential confounding effects related to different vocal capabilities and to maximize the number of trials available
for robust statistical analysis. Melodies outside this range, while part of the original experimental design, were
reserved for separate analyses addressing different research questions.
To align the vocal range of the singing stimuli with that of the participants, we assessed each participant’s vocal
capabilities. Male participants demonstrated an average vocal range of 28.5 semitones (range: 26–39), extend-
ing from a lower limit of C2–A2 to an upper limit of G4–C5. Female participants exhibited an average range of
35 semitones (range: 27–43), spanning from D3–A3 to B♭5–C7. Consequently, the selected tonal range for the
singing stimuli (B3–E♯4) fell well within the vocal capabilities of all participants (A3–G4). This careful matching
ensured that the fMRI data analysed reflected the neural responses to stimuli that participants could potentially
produce themselves, enhancing the relevance and interpretability of our findings related to the experience-
dependent neural activity during operatic voice perception.

Experimental procedure
Participants were positioned comfortably in the MRI scanner, with their heads secured to minimize movement.
They were instructed to remain still and focus on the auditory stimuli throughout the session. The fMRI experi-
ment was conducted over four functional runs using a block design, each approximately 12 min long. During
stimuli presentation, participants listened to a series of 18.25 s long melody blocks, containing four melodies of
the same singing style (either operatic or natural) and voice type (female-low, female-high, male-low, or male-
high). The melodies were played consecutively for approximately 4000 ms each, with an interstimulus interval
(ISI) of 750 ms between melodies. At the beginning of each run, there was a baseline period of 30 s. Following
the baseline, the melody blocks alternated with rest periods of varying durations (8, 11, or 14 s) to allow for
BOLD response decay and to reduce predictability of stimulus presentation. Each run included three blocks for
each stimulus type combination (style × voice type), resulting in 24 melody blocks and 25 baseline blocks per
run. During the experiment, each of the 48 melodies was presented once in a specific combination (e.g., melody
no. 1 in the natural style female-low condition). Each melody appeared twice in a run, but in different combina-
tions (e.g., melody no. 1 in the natural style female-low condition and in the operatic style male-high condition).
Auditory stimuli were delivered binaurally using MRI-compatible earphones (CONFON HP-IE-01, MR
Confon GmbH, Magdeburg, Germany). Visual fixation was maintained by a dark-grey dot displayed on a mid-
grey background on an MRI-compatible LCD screen (32″ NNL LCD Monitor, NordicNeuroLab, Bergen, Nor-
way; 1920 × 1080 pixels; 60 Hz). The screen was positioned at a distance of 142 cm from the participant’s eyes,
and viewed through a mirror attached to the MRI head coil. The presentation of stimuli and the timing of the
experimental procedure were controlled using custom-written scripts in MATLAB (R2015a, The MathWorks
Inc., Natick, MA, USA) utilizing the Psychophysics T ­ oolbox51,52.
To ensure participants’ attention to the auditory stimuli, they were engaged in a one-back memory task
during the melody blocks. Participants had to identify and remember if the current melody was identical to the
immediately preceding one and report the total number of one-back repetitions at the end of each run using a
response grip (NordicNeuroLab, Bergen, Norway).
Three additional runs featuring natural, operatic, and pure tone (sine wave) melodies were conducted in a
second fMRI session following a similar experimental setup. However, data from these additional runs were
reserved for separate analyses not included in the current study.

Data acquisition
Data were acquired on a Siemens Magnetom Prisma 3 T MRI scanner (Siemens Healthcare GmbH, Erlangen,
Germany) at the Brain Imaging Centre, Research Centre for Natural Sciences. A standard Siemens 32-channel

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head coil was applied. For functional measurements, a blipped-controlled aliasing in parallel imaging (CAIPI)
simultaneous multi-slice gradient-echo-EPI s­ equence53 was used with sixfold slice acceleration. Full brain cov-
erage was obtained with an isotropic 2 mm spatial resolution (208 × 208 mm field-of-view (FOV); anterior-to-
posterior phase encoding direction; 104 × 104 in-plane matrix size; 54 slices; 25% slice gap) and a repetition
time (TR) of 710 ms, without in-plane parallel imaging. A partial Fourier factor of 7/8 was used to achieve an
echo time (TE) of 30 ms. Flip angle (FA) was 59°. Image reconstruction was performed using the Slice-GRAPPA
­algorithm53 with LeakBlock ­kernel54. T1-weighted 3D MPRAGE anatomical imaging was performed using
twofold in-plane GRAPPA acceleration with isotropic 1 mm spatial resolution (TR/TE/FA = 2300 ms/3 ms/9°;
FOV = 256 × 256 mm).

Stimulus data processing

To assess the specific characteristics of vibrato in the singing stimuli (B3–E♯4) that fell well within the vocal
capabilities of all participants, fundamental frequencies ­(f0) of melodies matching participants’ vocal range were
extracted using the autocorrelation method in Praat (version 6.1.38). All ­f0 trajectories were segmented at tone
boundaries by thresholding the low-pass filtered ­(fc = 3 Hz) trajectories at the fundamental frequency of two
consecutive tones. To ensure clarity and remove transients, 50 ms were trimmed from the beginning and end of
each segment. Vibrato parameters, such as rate and extent, were determined using a discrete Fourier transform-
based ­method55 with refined spectral peak detection to accommodate tones lacking distinct vibrato. Parameters
were quantified by identifying the maximal spectral peak within the 4–8 Hz modulation range using Matlab’s
findpeaks() function. In cases where no spectral peak was detected, the vibrato rate was set to NaN, and the
vibrato extent was set to zero.

Processing of neuroimaging data

The functional images were first spatially realigned using SPM12’s two pass procedure. All realigned functional
images were then co-registered with the T1-weighted anatomical image, which also facilitated surface recon-
struction using the recon-all pipeline provided by FreeSurfer 7.1.156. Once co-registered, functional images
were resampled to fsaverage space and underwent iterative diffusion smoothing (40 iterations) as part of the
CONN preprocessing pipeline to increase signal-to-noise ratio and to compensate for intersubject variability
in functional activation.

Statistical modelling of neuroimaging data

For the first-level analysis of fMRI data, a standard voxel-wise general linear model (GLM) was implemented
using SPM12. This model included regressors for operatic and natural voice stimuli convolved with the canonical
hemodynamic response function to model BOLD responses. Only stimuli of the female-low and male-high voice
types (B3–E♯4 vocal range) were included as primary regressors for the purpose of this study, ensuring that the
analyses accurately reflected the participants’ physiological capabilities to produce these sounds. This selective
approach allows for a focused comparison of operatic versus natural singing styles within the participants’ own
vocal range, enhancing the relevance and interpretability of our findings. Additionally, operatic and natural
voice stimuli from the female-high (C5–F♯5) and male-low (G2–C♯3) ranges were also included as independent
regressors to account for their overall effects, although these data were not the focus of the current analysis and
are not discussed further.
Temporal high-pass filtering was applied with a cutoff frequency of 1/128 Hz to remove low-frequency drifts.
Movement-related variance was controlled by incorporating the estimated movement parameters from the rea-
lignment step, their temporal derivatives, and ­squares57 into the model as nuisance covariates. Additionally, these
movement-related parameters underwent band-stop filtering using a Butterworth IIR filter with a stopband of
0.2–0.5 Hz and a filter order of 12 to mitigate respiration-related ­effects58,59. Outlier scans, defined by framewise
displacement exceeding 0.9 mm or global BOLD signal changes surpassing 5 SDs in the Artifact Detection ­Tools60
implemented in the CONN toolbox were also accounted for using nuisance covariates. Temporal autocorrelations
were modelled using SPM12’s FAST model.
The first-level GLM produced contrast images based on parameter estimates (β values) of operatic and natural
stimuli regressors, which were then used as input for second-level whole-brain random-effects analyses (one-
sample t-tests). These primary analyses were conducted using a cluster-based permutation testing f­ ramework61
implemented in the CONN toolbox, applying a two-sided FDR-corrected P value threshold of P < 0.05 for cluster-
level significance to correct for multiple comparisons. Each significant cluster’s mass (sum of the t-squared
statistics across all voxels) and FDR-corrected P values (PFDR) are reported.
Preprocessing and statistical analyses of the imaging data were performed using the SPM12 (v7771) toolbox
(Wellcome Trust Centre for Neuroimaging, London, UK), the CONN (v.20.b) ­toolbox62, as well as custom-made
scripts running on MATLAB R2017a (The MathWorks Inc., Natick, MA, USA).

Region of interest (ROI) analysis

In our primary analysis above, we investigated the whole brain effect of stimuli style within the opera group
by comparing BOLD responses to operatic versus natural voice stimuli. Significant differences (see Results for
details) were observed bilaterally in early auditory regions, including the primary auditory cortex (A1), medial
belt (MBelt), posterior belt (PBelt), and lateral belt (LBelt) complexes, as well as in the A4 region of the auditory
association cortex, as identified by the Human Connectome Project Multi-Modal Parcellation atlas of human
cortical areas (HCP-MMP1.0)63. Additionally, significant effects related to stimuli style were detected bilaterally
in core areas of the default mode network (DMN), specifically the precuneus/posterior cingulate cortex (pCun/
PCC) and the medial prefrontal cortex (mPFC), according to the 200-region Kong2022 parcellation a­ tlas64.

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To determine if the effects observed in the voxel-wise analysis were unique to the opera group, we conducted
an ROI analysis across both groups using atlas-based ROIs. Percent signal change (PSC) of voxels was calcu-
lated for each participant within each ROI. Separate linear mixed effects (LME) models were then applied to
these PSC averages for the auditory and the core DMN ROIs. The models incorporated fixed-effects for group,
stimuli style, hemisphere, ROI, along with their interactions, and random effect for participant to account for
their inter-individual variability. Since stimuli style, hemisphere, and ROI are within-subject factors, and their
effect may vary significantly between individuals, we also included random slopes for these. Given the signifi-
cant age difference between groups (mean age was 39 in the opera and 29 in the control group), age could be a
confounding factor when investigating the group effects. To mitigate this, we controlled for age by including it
and its interaction with group as additional fixed effects.
Statistical analyses were performed in R (4.1.3)65 using the afex (1.2.1)66 and emmeans (1.8.5)67 packages.
Simple-effects were corrected for multiple comparisons using the single-step correction of the multcomp (1.4.23)
­package68.

Results
Vibrato characteristics of stimuli
Analysis of melodies within participants’ reproducible vocal range revealed distinct characteristics: the vibrato
extent was 16.5 ± 7.0 cents (mean ± SD) for natural tones and 126.5 ± 16.1 cents for operatic tones. Similarly,
vibrato rate was measured at 5.81 ± 0.57 Hz for natural and 5.76 ± 0.13 Hz for operatic tones. Melodies outside
this range, specifically from the male-low and female-high melody sets, were utilized for addressing different
research questions not covered in this report.

Whole brain analysis results

The whole brain analysis of the stimuli style contrast in opera singers revealed significant activations in sev-
eral brain regions involved in sound processing. Specifically, significantly higher BOLD responses were found
bilaterally for operatic compared to natural voice stimuli in the auditory cortex (left cluster mass = 74,484.6,
PFDR = 0.0042; right cluster mass = 50,768.28, PFDR = 0.0042), including the early auditory regions, such as the A1,
MBelt, PBelt, and LBelt, as well as the auditory association area A4. Furthermore, significant differences in BOLD
responses were found bilaterally, between operatic and natural voice stimuli in the core regions of the DMN,
namely in the pCun/PCC (left cluster mass = 17,346.09, PFDR = 0.013; right cluster mass = 9178.01, PFDR = 0.027)
and the mPFC (left cluster mass = 16,539.71, PFDR = 0.013; right cluster mass = 8336.37, PFDR = 0.027). Finally, we
identified a weak, yet statistically significant cluster (cluster mass = 5610.90, PFDR = 0.042) exclusively in the left
hemisphere, partially encompassing the left inferior parietal lobule (IPL) (see Fig. 1).
Conversely, the whole brain analysis by stimuli style in the control group (non-singers) revealed signifi-
cant activations confined to the bilateral auditory cortex. Notably, BOLD responses to operatic versus natu-
ral voice stimuli were significantly higher, manifesting bilaterally in the auditory cortex with nearly identical
regional extents to those observed in opera singers (left cluster mass = 82,857.32, PFDR = 0.00063; right cluster
mass = 84,947.41, PFDR = 0.00063).

ROI analysis results

To determine if the singing style effects identified in the opera group were specific to opera singers, we conducted
an atlas-based ROI analysis comparing BOLD responses between opera singers and controls across auditory
cortical and core DMN regions.

Auditory cortical regions

The ROI analysis targeting auditory cortical regions—specifically A1, MBelt, PBelt, LBelt, and A4—demonstrated
a significant main effect of stimuli style (F(1,63) = 77.47, P < 0.0001), indicating higher BOLD responses to oper-
atic versus natural voice stimuli across all ROIs and hemispheres for both opera singers and control participants.
Notably, the interaction between stimuli style and group was not significant (F(1,63) = 0.40, P = 0.53) (see Fig. 2),
highlighting a universal, experience-independent effect of stimuli style in the auditory cortex.

Core DMN regions

In the core DMN regions, specifically the pCun/PCC and the mPFC, our analysis revealed both a significant
main effect of stimuli style (F(1,63) = 8.26, P = 0.0055) and a significant interaction effect between group and
stimuli style (F(1,63) = 4.09, P = 0.047). This interaction was predominantly driven by a pronounced difference
in the opera group, where a significant preference for operatic over natural voice stimuli was observed (t = 3.44,
P = 0.0021); such a distinction was not evident in the control group (t = 0.61, P = 0.79). Specifically, opera singers
showed weaker negative BOLD responses to operatic compared to natural voice stimuli, a phenomenon not seen
in the controls (see Fig. 3). Additionally, no significant differences were found between groups when analysing
responses to operatic or natural stimuli independently (|t|≤ 1.64, P ≥ 0.16).

Discussion
In this study, we investigated how extensive musical training—in particular, Western operatic singing—can shape
brain responses to auditory perception of different singing styles. Focusing on vibrato, one of the acoustically
most salient features of operatic singing, we compared BOLD responses of trained opera singers and musically
untrained controls when listening to operatic (i.e., with vibrato) and natural (i.e., without vibrato) singing styles.
Whole-brain results demonstrated that opera singers exhibited increased BOLD responses in bilateral auditory

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Fig. 1.  Stimuli style effect on BOLD responses in opera singers. Heatmaps on inflated fsaverage cortical brain
models show second-level statistical t-maps of the operatic vs. natural stimuli style contrast thresholded at FDR
corrected P < 0.05. Significant stimuli style effects were revealed bilaterally in auditory cortical regions (A1,
MBelt, PBelt, LBelt, and A4) shown in lateral view (top), in core regions of the default mode network (pCun/
PCC and mPFC) shown in medial view (bottom) and only in the left hemisphere, partially overlapping with the
left IPL shown in lateral view (top left).

Fig. 2.  Stimuli style effect on BOLD responses in auditory cortical regions. Boxplots show the effect of stimuli
style (operatic vs. natural) and group (opera vs. control) on fMRI percent signal change (PSC) in both left (L)
and right (R) hemispheric (hemis) auditory regions (A1, MBelt, PBelt, LBelt and A4 ROIs).

cortical regions and weaker negative responses in core regions of the DMN—including the pCun/PCC and the
mPFC—as well as the left IPL when listening to operatic compared to natural singing voices. In contrast, among
non-singers, differential BOLD responses were restricted to the bilateral auditory cortex.
Building upon these findings, an ROI analysis highlighted significant group differences in BOLD responses
to operatic and natural singing voices. Importantly, an interaction effect within the DMN regions was driven by

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Fig. 3.  Stimuli style and group effects on BOLD responses in core default mode network (DMN) regions.
Boxplots show the effect of stimuli style (operatic vs. natural) and group (opera vs. control) on fMRI percent
signal change (PSC) in both left (L) and right (R) hemispheric (hemis) DMN regions (pCun/PCC and mPFC
ROIs).

a significant difference in the opera singers that was absent in controls. Specifically, opera singers showed weaker
negative BOLD responses to operatic compared to natural singing voices, suggesting that their extensive train-
ing and active experience with operatic singing have distinctly shaped neural responses to auditory perception
related to sound features inherent in their specialised singing style.

Whole brain results in opera singers and non‑singers

Differences in brain activation when comparing responses to operatic and natural singing voices were observed
in auditory cortical regions in both opera singers and controls, specifically, in bilateral early auditory cortical
regions and in the A4 complex of the auditory association cortex, a region associated with processing auditory
stimulus ­complexity69. These differences may be attributed to the amplitude and frequency-modulated vibrato
characteristic of operatic singing style. In addition to vibrato, a key feature of operatic singing is the spectral
envelope, which is harmonically more complex compared to natural singing. The phenomenon of vocal ring
in operatic voices, characterized by a clustering of frequency formants around three kHz, psychoacoustically
enhances perceived loudness. This acoustic feature allows trained singers to effectively project their voices over
the orchestral ­accompaniment70,71.
Previous studies have demonstrated that variations in acoustic properties can lead to increased activation
within the auditory cortex. For example, harmonic tones produce more activation than pure tones or single
frequencies in the right Heschl’s gyrus (HG) and bilaterally in the lateral supratemporal planes (STP)72. Simi-
larly, frequency-modulated tones elicit more activation than static tones in bilateral HG, the anterolateral part
of the lateral supratemporal planes, and the superior temporal s­ ulci72. The posterolateral STP, extending into
­ odulation73. These observations
the lateral auditory association cortex, is particularly responsive to frequency m
are consistent with the role of posterior non-primary auditory areas in processing spectrotemporally complex
­sounds74,75. Indeed, both amplitude (intensity) and frequency modulation trigger greater activations in these
regions compared to unmodulated t­ ones72. Given that the intensity of our singing-voice stimuli was normalized,
the primary distinctions between the stimuli lay in their spectrotemporal modulations. Therefore, the distinctive
acoustic properties of operatic singing voices likely account for the observed style-specific differences in auditory
cortex activation. This interpretation aligns with the lack of group difference in our ROI findings, indicating
that the enhanced auditory cortical responses to operatic singing were not solely linked to musical expertise.
However, only opera singers displayed style-specific activation patterns in cortical regions that are generally
associated with the DMN.

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ROI results
Building upon the voxel-wise findings, subsequent ROI analyses confirmed the absence of group differences in
the auditory cortex. However, a noticeable interaction effect between the group and singing style emerged within
the DMN, which was driven by the group of opera singers. Unlike the non-singer controls, opera singers exhibited
significant differences in BOLD responses between operatic and natural singing styles within core DMN regions,
particularly the mPFC and PCC. This result resonates with prior research demonstrating that music listening can
alter the DMN connectivity, depending on the level of self-referential processing the stimuli induce and prior
musical ­experience76,77, in addition to experience-dependent effects in neural networks associated with auditory
and sensorimotor, as well as cognitive and affective p ­ rocesses5,7,9,11.
The DMN plays critical roles in forming contextual associations, mediating emotional and self-referential
processes, fostering autobiographical memory, and facilitating mental state attribution, mind-wandering, and
­daydreaming78–80. According to the triple network model, network switching by the salience network alters the
dynamic temporal interactions between DMN and the frontotemporal ­networks81,82. Relevant sensory stimuli
can drive external event-driven switching from internally focused mental processes underscored by the DMN,
thus suppressing DMN and disrupting the internal narrative. The disengagement of the DMN from cognitive
control systems enables allocation of resources to serve external attention-demanding, goal-directed behaviours.
Through its characteristic connectivity and coactivation patterns, individual DMN nodes may support specific
network functions, as they are not uniformly engaged across cognitive domains. In this context, PCC and mPFC
are implicated as forming a core self-referential system within the DMN, with these nodes playing distinct roles
in differentiating self from o­ thers82.
Listening to music, especially when it evokes introspective thought and self-referential processes, has been
linked to increased DMN connectivity, as seen during aesthetic contemplation and engagements with preferred
­music76. This finding aligns with research showing that classical music training can rewire the DMN, leading to
enhanced connectivity within this network during resting-state ­fMRI83. Moreover, recent studies such as Mårup
et al.84 and Liao et al.85 have highlighted the DMN’s role in tasks that involve motor coordination and rhythm
production, particularly as these tasks become more automatic and internally driven in professional musicians,
consistent with the DMN’s function in maintaining internal representations and supporting cognitive processes
that require less focused attention during music ­production76,85,86.
However, in tasks involving free listening to complete musical pieces, the connectivity patterns in DMN-
related nodes appear differently for non-musicians and musicians. Non-musicians exhibit higher fMRI connectiv-
ity, suggesting introspective engagement, whereas musicians display altered patterns indicative of a more special-
ized auditory processing a­ pproach77. Similarly, using electroencephalography, decreased functional connectivity
between the ventromedial prefrontal cortex (vmPFC) and the pCun/PCC has been observed in musicians with
absolute pitch, as compared to non-musicians, when listening to whole orchestral ­works87. This divergence from
our findings could originate from the distinct acoustic stimuli used and their relevance to the specific sensori-
motor experiences of the musicians involved. Although speculative, it is possible that naturalistic music listen-
ing tasks engage musicians with diverse training backgrounds in more externally focused auditory processing
strategies, typically associated with DMN suppression. In contrast, the presence or absence of vibrato—a key
feature of operatic and natural singing styles—may consistently evoke autobiographical memories tied to internal
representations of embodied sensorimotor experiences of opera singers.
In line with this understanding, recent research has teased apart distinct conceptualizations of self-referential
processing within the DMN, highlighting its c­ omplexity79,88. For instance, activation of the vmPFC and dor-
somedial prefrontal cortex has been linked to valenced self-evaluation89. The activation of the frontal cluster,
which includes the vmPFC, corresponds to the degree of self-relevance of intero- or exteroceptive s­ timuli90.
Conversely, the posterior cluster, which encompasses the PCC, is thought to integrate self-referential informa-
tion into the temporal context and autobiographical m ­ emory91. These core regions of the DMN are thought to
facilitate the integration of incoming sensory inputs with prior internally generated representations to predict
and guide perception, learning, and action control in both naturalistic activities and cognitive tasks, suggesting
a role of the DMN in maintaining and updating detailed mental m ­ odels92,93. This enables experts to engage in
more efficient and automated cognitive p ­ rocesses94, as evidenced by our study observing increased DMN activ-
ity (reduced DMN deactivation) in trained opera singers when they listened to melodies sung in operatic style.
Thus, opera singers may integrate key auditory features of operatic singing with their extensive prior knowledge
and training. In combination with the increased engagement of auditory cortical areas, the DMN may aid in
more efficient processing and integration of training-specific auditory stimuli, highlighting the plasticity of the
DMN in adapting to the cognitive demands of expert-level singing ability.
This extensive and continuous learning process, necessary for mastering operatic singing, often requires
around 10,000 h of deliberate, goal-oriented practice before the age of 20 ­years95. Such rigorous training integrates
these complex skills into an individual’s self-model, impacting future self-perceptions and e­ valuations96,97. Our
findings demonstrate significantly higher activation within the vmPFC and PCC—key areas associated with
self-referential processing—when opera singers listened to vibrato, a hallmark of their trained style. This sug-
gests that the skills necessary to produce vibrato are deeply embedded in the singers’ self-models, influencing
their perception and engagement with operatically styled auditory cues. Notably, this heightened activation was
absent when opera singers listened to natural singing styles, and was not observed in the non-musician controls,
reinforcing the unique connection between specialized training and DMN activation.

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Conclusion
This study provides novel evidence suggesting that specialized musical sensorimotor training, as exemplified
by Western operatic singing, can significantly alter brain responses during the auditory perception of acoustic
features that are salient and relevant to one’s own singing style, such as vocal vibrato. These findings have pivotal
implications for understanding the neural underpinnings of musical training, particularly in the context of
vocal music, reinforcing the concept that our perceptions are actively shaped by past experiences and a­ ctions19.
The enhanced BOLD responses observed in the core DMN regions, specifically the vmPFC and the PCC, in
opera singers when listening to their own singing style underscore the DMN’s crucial role in self-referential
­processing82. These effects suggest that extensive training in operatic singing not only hones musical expertise but
also refines acoustically determined self-referential processes, potentially impacting an individual’s self-model.
This underscores the intriguing interplay between skill acquisition, self-representation, and neural plasticity.
Future research should delve deeper into this relationship, extending beyond cross-sectional designs to provide
more causal evidence of the effects of long-term singing training on cognition and behaviour.

Data availability
Data supporting the findings of this study are available from the corresponding authors on reasonable request.

Received: 18 September 2023; Accepted: 28 August 2024

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Acknowledgements
The current research was financially supported by the grants from the Hungarian Brain Research Program 3.0
to Z.V., the Hungarian Brain Research Program 2.0 under grant number 2017–1.2.1-NKP-2017-00002 provid-
ing support for J.Z., and the Danish National Research Foundation (DNRF117) and the Carlsberg Foundation
(CF22-1172) for B.K.

Author contributions
Conception and design: P.H., B.K., Z.V., J.Z.; Analysis and interpretation: A.B., P.H., Á.N., Z.V.; Data collection:
A.B., P.H., Á.N.; Writing the article: A.B., P.H., B.K., Á.N., Z.V., J.Z.

Competing interests
The authors declare no competing interests.

Additional information
Correspondence and requests for materials should be addressed to A.B. or Z.V.
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