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ABSTRACT Music possesses a remarkable capacity to evoke a broad spectrum of subjective responses and promote healing in humans, with rhythm playing a crucial role among its various elements.
Rhythmic entrainment is considered as one of the fundamental mechanisms, yet its subsequent behavioral responses and quantitative relationship remain unclear. In the study presented here,
we combined behavioral and electroencephalography experiments to explore the relationship between neural entrainment and emotional responses to rhythmic auditory stimuli, focusing on the
emotional dimensions of valence, arousal, and dominance. Our findings reveal that while all sequences across 12 different presenting rates significantly entrain neural oscillations,
sequences at different rates elicit distinct impacts on subjective emotional experience. The intensity of neural entrainment is associated with changes in emotional valence and dominance
under specific frequency conditions. This insight addresses a gap in the existing literature regarding the dominance dimension and provides potential for developing more targeted and precise
clinical interventions to enhance emotional well-being in the future. SIMILAR CONTENT BEING VIEWED BY OTHERS MUSICAL COMPONENTS IMPORTANT FOR THE MOZART K448 EFFECT IN EPILEPSY Article Open
access 16 September 2021 MUSICAL NEURODYNAMICS Article 18 March 2025 RELIABILITY FOR MUSIC-INDUCED HEART RATE SYNCHRONIZATION Article Open access 28 May 2024 INTRODUCTION The therapeutic
potential of music in modulating emotional states has been extensively acknowledged across both non-clinical and clinical settings, from alleviating anxiety in cancer patients1 to improving
mood regulation in individuals with neurodegenerative disorders2. Notably, rhythmic auditory stimulation, when devoid of the melodic complexity inherent in music, demonstrates comparable
efficacy in enhancing emotional well-being3,4. Various forms of rhythmic auditory stimulation have been reported to alleviate emotional symptoms. For instance, a study demonstrated that the
Schumann resonance frequency (7.83 Hz), a special frequency between the theta and alpha bands, when presented as an isochronous tonal sequence, effectively reduced anxiety and promoted
emotional calmness5. Furthermore, patients with mild anxiety exhibited significant improvements in anxiety symptoms following exposure to a mixture of theta and delta binaural beats6. While
the neural mechanisms of rhythmic stimulation are well-established in motor rehabilitation, language therapy, and attention regulation7, how the structured rhythmic patterns confer emotional
benefits remains obscured. Emotional processing fundamentally relies on distributed neural networks8 where oscillatory activity coordinates cross-regional communication9,10. Crucially,
previous studies have shown that specific neural oscillation features are correlated with the key dimensions of emotion: valence (a bipolar continuum from displeasure to pleasure), arousal
(spanning sleep to frenzied excitement), and dominance (ranging from feelings of helplessness to control), as conceptualized in the three-dimensional emotion model11. For instance, high
arousal stimuli have been linked to a decrease in alpha and beta band power12, whereas alpha asymmetry in parieto-occipital networks correlates with emotional valence processing13.
Dominance, though less studied, may engage beta oscillations in frontoparietal control networks14. Overall, low frequency oscillations of delta and theta ranges are associated with
motivational and emotional processes15. Beyond emotion detection and processing, oscillatory activity also plays a crucial role in emotion regulation. Increased frontal theta power is
associated with successful cognitive reappraisal16, and theta oscillations over frontal-midline regions have been robustly associated with emotional arousal regulation17, establishing a
physiological link between brain oscillations and emotion regulation processes. These frequency-specific dynamics create “oscillatory gateways”, where external rhythmic stimuli that match
endogenous brain frequencies could selectively modulate emotional dimensions18,19. Importantly, these frequency-specific oscillations are also involved in music processing. Rhythmic external
stimuli can induce phase locking and enhance the neural oscillatory activity of corresponding frequency bands in the brain. This neural entrainment, defined as the stimulus-driven
synchronization of neuronal oscillations to periodic external inputs19,20, typically manifests through two complementary neurophysiological signatures21: phase coherence, which refers to the
consistency of phase alignment across stimulus repetitions, reflecting temporal precision of entrainment22, and stimulus-evoked power enhancement, wherein amplitude amplification at the
driving frequency, indicating resonance strength23. Previous research has demonstrated various forms of auditory rhythms, such as isochronous sound sequences, monaural or binaural rhythms,
and music, can induce entrainment, though the effects vary24. Both monaural and binaural beats in the theta frequency band can modulate subcortical response and cortical activity, with
monaural beats showing stronger modulation25. Research utilizing human intracranial electroencephalography has observed that a 5 Hz binaural beat increases temporal phase synchronization,
while a 5 Hz monaural beat leads to decreased mid-temporal synchronization26. The synchronization theory between neural oscillations and external rhythms has been validated also in studies
using real music beats as stimuli, where beat frequency and its higher-order harmonics modulate neural responses27,28. Beat frequencies selectively enhance EEG power through meter-specific
synchronization29, demonstrating the brain’s preferential response to rhythmically structured inputs. However, despite these advancements, the precise mechanisms by which rhythmic auditory
stimuli—particularly those with frequency-specific characteristics—affect emotional states remain a subject of ongoing debate. These unresolved questions serve as the impetus for our
investigation into how different rhythmic frequencies influence emotion, with a particular focus on neural entrainment and emotional experience. Building on the above framework, we
hypothesize that musical rhythm acts as an external “oscillatory pacemaker”, entraining emotion-related circuits through frequency-matched phase alignment. Hence, in order to contribute more
evidence in this field and investigate the entrainment of neural oscillations to rhythmic auditory stimuli, specifically examining how tonal sequences of varying frequencies affect
subjective emotional experiences, we conducted a combined behavioral and EEG experiment in which participants were required to listen to twelve tonal sequences with different frequencies and
to assess their affective responses through self-reported valence, arousal and dominance ratings. We hypothesize that rhythmic tonal sequences will induce frequency-specific neural
entrainment corresponding to their stimulation rates, modulate emotional dimensions (valence, arousal, dominance) differently through entrainment-mediated oscillatory changes by exhibiting
relationships between entrainment strength and emotional response magnitude. Our research aims to elucidate the mechanisms by which rhythmic auditory input influences affective states,
focusing on the relationship between neural entrainment and emotional responses. In addition, this study also seeks to provide potential implications and inspiration for the clinical
therapeutic use of music in mental disorders, emotional troubles and cognitive training or rehabilitation by offering a deeper understanding of potential mechanisms. MATERIALS AND METHODS
PARTICIPANTS A total of 40 graduate students were recruited for this study from colleges and universities. Due to the technical issues, 3 participants’ recordings were invalid, who were
excluded from the data analysis, ending up with 37 participants (19 males and 18 females, mean age = 22.72 ± 1.91 years) with balanced gender and age. All participants had normal hearing and
either normal or corrected-to-normal vision, with no reported history of mental disorders or severe physical illnesses. The study was conducted in accordance with the Declaration of
Helsinki and approved by the local ethics committee (SMHC-IRB: 2021-45). Participants gave their written informed consent and received monetary compensation for their participation. STIMULUS
AND PROCEDURE The experiments were conducted in a regular room. Acoustic stimuli were delivered to the participant via a wired loudspeaker (BMS09, Lenovo) through a PC (171501-AQ, Xiaomi).
Stimuli were delivered at a comfortable sound level (65 ~ 75 dB) adjusted according to each participant’s report. Testing protocols were executed with a custom MATLAB (MathWorks, Natick, MA)
program developed in our laboratory. Tonal sequences were composed of eight musical notes from the F major scale (F, G, A, Bb, B, C, D and E). To ensure the tonal sequences were musically
structured as defined by the Western tonal system, a transition matrix providing the transition probabilities between pitch classes was adopted30. Pitch classes – that is, pitch regardless
of octave – form the state space of the Markov chain, while the relative frequencies with which one pitch class leads to the following pitch class form the matrix of transition
probabilities. To ensure the naturalness of the sound of the musical note, the sound file of each note was generated by Noteflight31. The sound files were imported into MATLAB and 12 tonal
sequences were generated based on the transition matrix. The sequences obeyed to 1 to 12 Hz as the presenting rate of the tonal notes (Fig. 1B). That is for 1 Hz stimulus, one note was
presented per second; for 2 Hz stimulus, two notes were presented per second, etc. Each sequence lasted for one minute. The selection of the particular frequency range (1–12 Hz) was guided
by three key considerations from neuroscientific and music cognition perspectives: (1) As stated in the introduction, the 1–12 Hz spectrum encompasses major oscillatory bands implicated in
distinct emotional processes. This coverage allows systematic investigation of frequency-specific emotion-circuit engagement; (2) Ecological validity in musical rhythm perception: While
beta/gamma oscillations (> 12 Hz) are relevant for higher-order cognitive processing, musical beat perception exhibits strict temporal constraints. As demonstrated by London32, the human
pulse perception window spans 0.5–10 Hz (30–600 bpm), with upper bounds constrained by auditory temporal resolution. Our 12 Hz upper limit slightly exceeds this range to ensure full coverage
of musically-relevant tempi while avoiding non-perceptual pulse rates (> 12 Hz ≈ 720 bpm); (3) Addressing methodological limitations in prior work: Existing studies typically employ
sparse frequency sampling (e.g., comparing only fast vs. slow tempo), potentially obscuring nuanced frequency-emotion relationships. Our 1–12 Hz continuum with 1 Hz resolution provides
capacity to disentangle overlapping functions within broad bands. All the participants listened to the same 12 sequences but with different orders (Fig. 1A). The 12 sequences were randomly
presented to participants. The spectrum of the envelope of the 12 stimuli was showed in Fig. 1C. The variance in emotional assessments, before and after listening to each piece, were
accounted for by three major dimensions: affective valence, arousal, and dominance33, by a series of Self-Assessment Manikins (SAM)34 Scales using 9-point scales (1–9). EEG DATA COLLECTION
AND PREPROCESSING The EEG was continuously recorded from 64 Ag/AgCl electrodes using a NeuSenW amplifier (Neuracle, China). Electrode positions conformed to the extended 10–20 system. All
channels were digitized at a sampling rate of 1000 Hz. AFz served as the ground and CPz as the online reference. Electrode impedances were below 10 kΩ at the beginning of the experiment. EEG
data were preprocessed offline using a combination of the EEGLAB (version 14.1.2b), and FieldTrip (version 20200130) toolboxes in MATLAB (version 2019b; MathWorks), as well as
custom-written MATLAB scripts. Continuous data from each run were imported into EEGLAB and the non-EEG channels were excluded. The data were then high-pass filtered (pass-band edge = 0.5 Hz,
transition width = 0.5 Hz) using a zero-phase, hamming-windowed, finite impulse response (FIR) sinc filter (implemented in the pop_eegfltnew function from the frflt plugin; v2.4), followed
by a low-pass filter with the stop frequency being 120 Hz. Data were then filtered for line noise using a band-stop filter at 50 and 100 Hz (0.5 Hz bandwidth). The flatline and cross-talk
channels were identified and spherically interpolated, which were processed with artifact subspace reconstruction (implemented with the pop_clean_artifacts function from the clean_rawdata
plugin; v2.3 with flatline criterion = 5 s; channel correlation criterion = 0.80). Data were then re-referenced to the common average reference (CAR) prior to independent component analysis
(binica implementation of the extended logistic infomax ICA algorithm). Resultant decompositions were automatically classified using the ICLabel plugin (v1.342). Components classed as either
ocular or cardiac in origin with > 90% probability were subtracted. The ICA weight matrix was subsequently applied to the original data. Note that the maximum number of ICA components to
be extracted was determined by the residual degrees of freedom after SSS rank reduction during MaxFiltering. Continuous data were then segmented in epochs of 2 s with no overlap, as the
lowest stimulus frequency was 1 Hz and 2 s contained two complete circles for further analyses. The epochs with a maximum amplitude of any channel larger than 100 uV were discarded from
further analyses. 31 participants with more than 90% valid trials were included for the further analysis, which resulted in a total rejection of 1.33% of all trials. After these steps, data
were exported from EEGLAB into the FieldTrip data format for further analyses. EXTRACTING ENTRAINMENT FEATURES We employed two distinct methods to measure the entrainment of rhythmic tonal
sequences on neural oscillations: spectral measurement and phase measurement. Specifically, we focused on evoked responses for spectral analysis and inter-trial phase coherence (ITPC) for
phase analysis. To extract these entrainment features, each 2 s epoch was taken as a trial. The EEG response in each trial was converted into the frequency domain of 1–15 Hz using the DFT
with a 0.5 Hz step. The complex-valued Fourier coefficient of a certain frequency bin was summed across trials, and evoked power spectrum (EPS) of this frequency is the normalized squared
magnitude of the sum by dividing the trial number. It is the same as the power spectrum of the EEG response waveform averaged over trials. The EPS reflects the power of EEG responses that
are synchronized to the tonal sequence. The ITPC reflects the consistency of phase angles across multiple trials at a specific frequency, which measures how aligned or synchronized the
phases of a particular frequency component are across different trials. The stimulus-driven phase alignment of endogenous oscillations, quantified through ITPC at the stimulation frequency,
could then reflect neural entrainment. First, the complex Fourier coefficients are normalized by their magnitude to isolate the phase information, resulting in complex numbers with unit
magnitude. These normalized complex values are then summed across trials for each time-frequency point. The absolute value of this sum is taken and normalized by the number of trials,
yielding a value between 0 and 1, where 1 indicates perfect phase coherence across trials and 0 indicates no coherence. STATISTICAL ANALYSIS The statistical significance of the neural
response at a target frequency was tested for EPS and ITPC, respectively. For both features, we conducted repeated measures analyses of variance (ANOVAs) to examine the effects of tonal
presenting rate and the response type (target vs. non-target). The target frequency was defined as matching the tonal presenting rate, while non-target frequencies were neither the tonal
presenting rate nor its harmonics. If Mauchly’s test indicated that the assumption of sphericity is violated, Greenhouse-Geisser correction was adopted for the factors with more than 2
levels. In cases where significant interactions were observed, post hoc tests were conducted using false discovery rate (FDR) corrected pairwise comparisons to identify specific differences
between target and non-target for each tonal presenting rate. Behavioral data were also analyzed using repeated measures ANOVAs. The influence of sampling time (before or after listening)
and the tonal presenting rate on the three emotional ratings was examined. Similarly, if Mauchly’s test indicated that the assumption of sphericity is violated, Greenhouse-Geisser correction
was adopted for the factors with more than 2 levels. In cases where significant interactions were observed, post hoc tests were conducted using FDR corrected pairwise comparisons to
identify specific differences between before and after listening for each tonal presenting rate. Finally, for the frequency which demonstrated a significant emotional influence, a
correlation analysis was conducted to explore the relationship between the modulation intensity of neural entrainment at the target frequency and the corresponding emotional changes. Pearson
correlation coefficients were computed between entrainment magnitude (averaged across electrodes) and standardized emotion change scores (post- vs. pre-task). The significance level for all
statistical tests was set at α = 0.05. Effect sizes were calculated using partial eta-squared (η²) for ANOVA results and Cohen’s d for post hoc comparisons. RESULTS ENTRAINMENT OF RHYTHMIC
TONAL SEQUENCE ON NEURAL OSCILLATION We first analyzed the global field power of EEG responses (Fig. 2A and D). In this analysis, the ITPC and EPS were calculated for each electrode and then
averaged over electrodes. In the grand average over subjects, the response to each frequency showed clear peaks at the target rates and their harmonics for both phase (Fig. 2A) and spectral
(Fig. 2D) features, respectively. The results of the repeated measures ANOVAs demonstrated significant interaction between tonal presenting rate and the response type for both ITPC [F(11,
330) = 4.81, _p_ < 0.001, η2 = 0.14] and EPS [F(11, 330) = 14.10, _p_ < 0.001, η2 = 0.32]. While further comparison between response to target frequency and non-target frequency of
global response showed that, for all the 12 presenting rates, the neural responses at target frequencies were significantly larger than those at non-target frequencies (Table 1) for both
ITPC (Fig. 2B) and EPS (Fig. 2E). To further elucidate the spatial distribution of entrainment effects, we examined the topographical patterns of EEG activity across the scalp. From the
topography, we observed that the rhythmic sequences of 12 different rates significantly modulate neural oscillatory activity at corresponding frequencies, with a particularly pronounced
effect in the frontocentral area, which is consistent between ITPC (Fig. 2C) and ESP (Fig. 2F). THE IMPACT OF RHYTHMIC TONAL SEQUENCE ON SUBJECTIVE EMOTION As shown in Fig. 3A, the averaged
changes in subjective emotions of three dimensions after listening to the tonal sequences were influenced differently compared with before listening. In particular, all tonal sequences below
6 Hz decreased participants’ valence feelings while increasing their dominance feelings (Table 2). Conversely, sequences above 6 Hz increased participants’ valence feelings (except for 10
Hz, FDR corrected _p_ = 0.827) while decreasing their dominance feelings. While for arousal, the changes were turbulent (Table 2). The statistical analyses along the 12 presenting rates
revealed that the dominance dimension was significantly affected by the interaction between time and frequency [F(11,396) = 3.20, _p_ < 0.001, η2 = 0.08] and marginally significantly
affected by the time factor [F(1,36) = 3.88, _p_ = 0.057, η2 = 0.10]. However, the valence and arousal dimensions did not show significant effects for any of the factors (Table 2). Post hoc
analysis with FDR correction further investigated the significant interaction effect observed in the dominance dimension. The results showed a significant impact of the tonal sequence at 3
Hz (FDR corrected _p_ = 0.032) and 10 Hz (FDR corrected _p_ = 0.031), which indicated that 3 Hz and 10 Hz could notably change the level of emotional dominance of participants. No
significant differences were found for the valence and arousal dimensions at any specific rates (Table 2). THE RELATIONSHIP BETWEEN NEURAL ENTRAINMENT AND THE CHANGES OF EMOTION At last, the
relationship between the modulation intensity of neural entrainment, in terms of ITPC and ESP, and emotional changes was tested. Among all the conditions showing at least marginal
significance in the behavioral data analysis, the neural entrainment of the tonal sequences with presenting rates above 6 Hz demonstrated a significantly positive correlation between
modulation intensity of ITPC and the absolute changes of emotional valence (_r_ = 0.22, _p_ = 0.002) and dominance (_r_ = 0.16, _p_ = 0.030) (Fig. 3B). That’s, as the rhythmic tonal
sequences with high presenting rates made participants feel more happy but less dominant of self’s emotion, the stronger the modulation intensity is the stronger emotional influence the
rhythmic tonal sequences could make. However, the neural entrainment of tonal sequences with presenting rates below 6 Hz showed a weak or negligible correlation between ITPC and absolute
changes in valence (_r_ = 0.00, _p_ = 0.969) and dominance (_r_ = -0.05, _p_ = 0.549) (Fig. 3B). DISCUSSION The present work integrates electrophysiological recordings and subjective
emotional assessments to explore the link between neural entrainment and emotional reactions to rhythmic auditory input. Although all the rhythmic tonal sequences of 12 presenting rates can
significantly entrain the neural oscillation, sequences at varying rates have distinct effects on the subjective emotional states. Furthermore, under the specific frequency stimuli, the
strength of neural entrainment correlates with changes in emotional valence and dominance. This approach addresses three limitations of prior research: First, complex musical stimuli often
conflate rhythmic and melodic effects35; Second, more comprehensive assessment of emotional changes including the neglected dominance dimension is conducted; Third, links between entrainment
magnitude and emotional changes are quantified. Music is known to induce strong affective reactions in listeners both in the form of pleasure and emotional responses. However, the
psychological mechanisms that drive these processes are still poorly understood. Entrainment is the physical principle whereby temporally organized systems, or oscillators, tend to become
synchronized as they interact with each other. Given the ubiquity of rhythmical patterns in both music and humans, rhythmic entrainment has been proposed as a potential emotion induction
mechanism during music listening. The analysis results from our experiments provide deep insights into how different presenting rates with varying frequencies influence neural responses, as
evidenced by both EPS and ITPC measures (Fig. 2). The choice of EPS and ITPC as indicators of neural entrainment aligned with two characteristics of entrainment at the population level, i.e.
phase alignment and amplitude-increase19. The clear peaks observed in the global field power of EEG response of EPS and ITPC at target frequencies and the harmonics suggest that the brain’s
neural ensembles can robustly synchronize to those external tonal stimuli (Fig. 2A and D). This synchronization is not merely a general phenomenon but appears to be strongly
frequency-specific, as indicated by the significantly larger neural responses at target frequencies compared to non-target frequencies across all 12 presenting rates (Fig. 2B and E). Other
study also found similar results. Nozaradan and colleagues found that the magnitude of the SS-EPs elicited at frequencies related to beat and meter perception was significantly higher
compared to the SS-EPs elicited at frequencies not associated with beat and meter36. Furthermore, our results aligned with the auditory steady-state response, which describes the phenomenon
where neural ensembles exhibit an increase in power amplitude at the frequency of the auditory stimulation37. Previous researchers using intracranial electroencephalography (iEEG) recordings
also demonstrated that auditory steady-state responses can occur at various frequencies (4–16 Hz) across different regions of the primary auditory cortex38. The spatial distribution of
entrainment effects is shown in Fig. 2C and F. The topography revealed that this neural entrainment effect is not uniformly distributed across the whole scalp but is particularly pronounced
in the frontocentral area. These patterns align closely with the scalp topographies observed in previous EEG measurements of entrainment responses to the acoustic envelope, which are
characterized by a prominent fronto-central to posterior dipole21,36,39. The low-frequency BOLD signals that characterize the frontoparietal network are linked to relatively slow brain
oscillations, particularly in the θ or α frequency range detectable through electrophysiological recordings. This connection likely plays a crucial role in facilitating the coordination of
activities across the entire brain network40. Previous research has often directed attention to dichotomous categories of musical emotions, such as pleasant emotions against unpleasant
emotions or emotions arranged along the same two fundamental dimensions of valence and arousal as other basic emotions41. However, the emotional dominance has been insufficiently addressed
for an extended period. Previous literature has also highlighted the potential and the often overlooked importance of the dominance component in contextualized music listening42. Dominance
refers to the feelings of control or influence on events or surroundings to what extent, which makes it possible to distinguish angry from anxious, alert from surprised, relaxed from
protected, and disdainful from impotent11. Based on the data-driven analysis, a boundary emerged. Tonal sequences below 6 Hz heightened participants’ feelings of dominance, while sequences
above 6 Hz diminished these feelings. Furthermore, among the twelve stimulating rates tested, 3 Hz and 10 Hz could particularly alter participants’ levels of emotional dominance, which may
have unique implications for quantitative manipulation for precise intervention of emotional regulation in therapeutic interventions. While this division was based on data-driven discovery,
its neurobiological plausibility can be supported by two key observations: (1) Theta-alpha transition zone: 6 Hz resides at the theoretical interface between theta (4–8 Hz) and low-alpha
(8–10 Hz) bands. Recent work identifies 5–7 Hz as a functional transition point where sensory-driven theta rhythms shift to cognitive alpha oscillations43. (2) Cortico-subcortical resonance
split: Aligns with hippocampal-cortical theta (3–6 Hz) crucial for emotional memory consolidation44 and 6 Hz corresponds to thalamocortical alpha generation (8–10 Hz) involved in affective
filtering45. By contrast, our results indicated that valence alteration has the opposite pattern compared to the change of dominance. All tonal sequences below 6 Hz decreased participants’
valence feelings while sequences above 6 Hz increased their valence feelings. The former part may align with the fact that theta (4–8 Hz) has been shown to increase negative affect in some
circumstances46. However, the previous research related to the application of auditory beat stimulation manipulating cognitive processes or modulating affective states yielded contradictory
results3. Meanwhile, the effectiveness of beat stimulation can be influenced by various factors, including the length of time the stimulus is applied. For instance, quite a few researches
have pointed out that exposure to the delta range (0–4 Hz) binaural beat would reduce the anxiety level6,47,48, conforming to our findings since when anxiety decreases, individuals typically
feel a greater sense of control over the situation, leading to a corresponding increase in the dominance dimension. Nevertheless, among these effective treatments mentioned above, except
for one intervention lasting only thirty minutes, the remaining extended from one month to sixty days, and maybe this is the reason why their findings are inconsistent with our results of
stimuli below 6 Hz decreasing valence. What’s more, patients who received 10 Hz binaural-beat stimulation for 20 minutes showed a notable reduction in anxiety scores after the intervention
compared to those who did not receive the stimulation49, which means a rise in the emotional dominance. Given the divergence between our findings and those of Weiland, further research is
needed to explore the underlying factors contributing to these differences. Interestingly, we found that the average changes in emotional arousal are turbulent and do not show significant
effects for time factor and the interaction between time and frequency. It aligned with a previous study in which found no variations in heart rate or skin conductance, utilized as indicator
of emotional arousal, were detected across any of the beat frequencies used50. The lack of clear patterns could suggest that while rhythmic stimulation impacts other dimensions of emotion,
like valence or dominance, its effect on arousal might be less straightforward, potentially requiring more nuanced or longer-term studies to fully capture its dynamics. Further exploration
into these factors could provide deeper insights into how and when rhythmic sequences effectively modulate emotional arousal. In the final part, we try to study in detail the affective
consequences of neural modulations, thereby enhancing the understanding of relationship between modulation intensity and emotion alteration. We found that the neural entrainment induced by
tonal sequences with presentation rates exceeding 6 Hz showed a significant positive correlation between the modulation intensity of ITPC and the absolute changes in emotional valence and
dominance. In other words, as the high-rate rhythmic tonal sequences made participants feel happier but less in control of their emotions, the greater the modulation intensity, the more
pronounced the emotional impact of the rhythmic sequences. This suggests that the degree of neural entrainment significantly influences emotional valence and dominance, providing evidence
that rhythmic entrainment, specifically in the neural level, may be one of the underlying mechanisms through which music evokes emotions. The observed correlation between entrainment
magnitude and emotional intensity may arise from two distinct yet complementary pathways. In the interoceptive channel, theta-band entrainment modulates autonomic resonance via vagal nerve
regulation, generating visceral feedback that anchors emotional qualia in the body51. Simultaneously, even passive listening to musical rhythms recruits motor regions of brain52, and neural
synchronization in the motor channel could engage cortico-striatal circuits involved in predictive movement planning, generating approach-avoidance motivational states that translate
rhythmic patterns into embodied affect53. Such sensorimotor predictions may simulate emotional resonance through corticospinal coherence with auditory rhythms, as posited by embodied
simulation theories. These findings underscore the complex relationship between neural oscillations and emotional experiences, suggesting that different frequency ranges might differentially
impact affective states. The proposed framework represents a testable hypothesis, rather than an established mechanism, warranting further investigation. Understanding these nuanced
interactions could have implications for developing therapeutic interventions that utilize rhythmic auditory stimuli to modulate emotional states, particularly in clinical settings where
managing affective disorders is crucial. Future research should explore the underlying neural mechanisms more deeply and consider individual differences in susceptibility to these
frequency-specific effects. While these findings provide valuable insights into the relationship between neural oscillations and emotional experiences, it’s important to acknowledge the
limitations of the current study and consider directions for future research. A key limitation is the relatively small sample size, which may affect the generalizability of the findings.
Additionally, the specific frequencies selected and the duration of the stimuli may have influenced the outcomes, potentially limiting the broader applicability of the results. Previous
research by Thut et al.19 has reported that sustained entrainment effects are likely short-term, ceasing shortly after the end of the stimulus. This observation raises questions about the
duration for which the evoked emotions are retained and highlights the need for further investigation into the temporal dynamics of these effects. Understanding this could have significant
implications for designing clinical interventions that aim to induce or sustain particular emotional states. Except for further considering the duration of stimulation mentioned above,
future research should explore a broader range of rates in rhythmic stimulation to better understand how different stimulating rates influence neural entrainment and emotional responses.
This could identify specific ranges that are particularly effective for mood and cognitive interventions, leading to more targeted and efficient intervention. Additionally, using advanced
brain imaging techniques like fMRI or MEG could offer deeper insights into the neural mechanisms of neural entrainment. These tools would help visualize the brain regions and networks
involved, paving the way for developing more precise and personalized therapeutic approaches. CONCLUSION This study employed a combination of electrophysiological measurements and subjective
emotional evaluations to investigate the relationship between neural entrainment and subjective emotional responses triggered by the rhythmic tonal stimuli. Across the 12 presentation rates
of rhythmic tonal sequences examined, sequences at varying frequencies influenced the subjective emotion in varying ways. Notably, it sheds light on the emotional dimension of dominance, an
area often underappreciated in previous studies. As all the sequences significantly entrain the neural oscillation, the nuanced relationship between neural entrainment and subjective
emotional responses warrants further exploration. Expanding on these aspects could provide deeper insights into the neural mechanisms underlying emotional processing and contribute to more
effective clinical applications for emotional health. DATA AVAILABILITY Data is provided within the manuscript or supplementary information files. ABBREVIATIONS * EEG: Electroencephalography
* EPS: Evoked power spectrum * ITPC: Inter-trial phase coherence REFERENCES * Burns, D. S. The effect of the Bonny method of guided imagery and music on the mood and life quality of Cancer
patients. _J. Music Ther._ 38 (1), 51–65. https://doi.org/10.1093/jmt/38.1.51 (2001). Article CAS PubMed Google Scholar * Raglio, A. Effects of music and music therapy on mood in
neurological patients. _World J. Psychiatry_. 5 (1), 68. https://doi.org/10.5498/wjp.v5.i1.68 (2015). Article PubMed PubMed Central Google Scholar * Chaieb, L., Wilpert, E. C., Reber, T.
P. & Fell, J. Auditory beat stimulation and its effects on cognition and mood States. _Front. Psychiatry_. 6 https://doi.org/10.3389/fpsyt.2015.00070 (2015). Article PubMed PubMed
Central Google Scholar * Chee, Z. J. et al. The effects of music and auditory stimulation on autonomic arousal, cognition and attention: A systematic review. _Int. J. Psychophysiol._ 199,
112328. https://doi.org/10.1016/j.ijpsycho.2024.112328 (2024). Article PubMed Google Scholar * Ray, R. W. Isochronic Tones in the Schumann Resonance Frequency for the Treatment of
Anxiety: A Descriptive Exploratory Study (Order No. 10618726). Available from ProQuest Dissertations & Theses Global. (1980860439). (2017).
https://www.proquest.com/dissertations-theses/isochronic-tones-schumann-resonance-frequency/docview/1980860439/se-2 * Scouranec, R. P. L. et al. &. Use of binaural beat tapes for
treatment of anxiety: A pilot study of tape preference and outcomes. _Altern. Ther. Health Med._ 7 (1), 58–63. (2001). * Thaut, M. & Hoemberg, V. _Handbook of Neurologic Music Therapy_
(Oxford University Press, 2014). * Malezieux, M., Klein, A. S. & Gogolla, N. Neural circuits for emotion. _Annu. Rev. Neurosci._ 46 (1), 211–231.
https://doi.org/10.1146/annurev-neuro-111020-103314 (2023). Article CAS PubMed Google Scholar * Buzsáki, G. & Draguhn, A. Neuronal oscillations in cortical networks. _Science_ 304
(5679), 1926–1929. https://doi.org/10.1126/science.1099745 (2004). Article ADS CAS PubMed Google Scholar * Knyazev, G. G. EEG delta oscillations as a correlate of basic homeostatic and
motivational processes. _Neurosci. Biobehav. Rev_. 36 (1), 677–695. https://doi.org/10.1016/j.neubiorev.2011.10.002 (2012). Article Google Scholar * Russell, J. A. & Mehrabian, A.
Evidence for a three-factor theory of emotions. _J. Res. Pers._ 11 (3), 273–294. https://doi.org/10.1016/0092-6566(77)90037-X (1977). Article Google Scholar * Schubring, D. & Schupp,
H. T. Emotion and brain oscillations: High arousal is associated with decreases in Alpha- and lower Beta-Band power. _Cereb. Cortex_. 31 (3), 1597–1608.
https://doi.org/10.1093/cercor/bhaa312 (2021). Article PubMed Google Scholar * Harmon-Jones, E., Gable, P. A. & Peterson, C. K. The role of asymmetric frontal cortical activity in
emotion-related phenomena: A review and update. _Biol. Psychol._ 84 (3), 451–462. https://doi.org/10.1016/j.biopsycho.2009.08.010 (2010). Article PubMed Google Scholar * Lindquist, K. A.,
Satpute, A. B., Wager, T. D., Weber, J. & Barrett, L. F. The brain basis of positive and negative affect: Evidence from a Meta-Analysis of the human neuroimaging literature. _Cereb.
Cortex_. 26 (5), 1910–1922. https://doi.org/10.1093/cercor/bhv001 (2016). Article PubMed Google Scholar * Harmony, T. The functional significance of delta oscillations in cognitive
processing. _Front. Integr. Nuerosci._ https://doi.org/10.3389/fnint.2013.00083 (2013). 7. Article Google Scholar * Ertl, M., Hildebrandt, M., Ourina, K., Leicht, G. & Mulert, C.
Emotion regulation by cognitive reappraisal—the role of frontal theta oscillations. _NeuroImage_ 81, 412–421. https://doi.org/10.1016/j.neuroimage.2013.05.044 (2013). Article PubMed Google
Scholar * Aftanas, L. I. & Golocheikine, S. A. Human anterior and frontal midline theta and lower alpha reflect emotionally positive state and internalized attention: High-resolution
EEG investigation of meditation. _Neurosci. Lett._ 310 (1), 57–60. https://doi.org/10.1016/S0304-3940(01)02094-8 (2001). Article CAS PubMed Google Scholar * Fries, P. Rhythms for
cognition: Communication through coherence. _Neuron_ 88 (1), 220–235. https://doi.org/10.1016/j.neuron.2015.09.034 (2015). Article CAS PubMed PubMed Central Google Scholar * Thut, G.,
Schyns, P. & Gross, J. Entrainment of perceptually relevant brain oscillations by Non-Invasive rhythmic stimulation of the human brain. _Front. Psychol._ 2, 170.
https://doi.org/10.3389/fpsyg.2011.00170 (2011). Article PubMed PubMed Central Google Scholar * Zoefel, B., ten Oever, S. & Sack, A. T. The involvement of endogenous neural
oscillations in the processing of rhythmic input: More than a regular repetition of evoked neural responses. _Front. NeuroSci._ 12 https://doi.org/10.3389/fnins.2018.00095 (2018). * Ding, N.
et al. Characterizing neural entrainment to hierarchical linguistic units using electroencephalography (EEG). _Front. Hum. Neurosci._ 11. https://doi.org/10.3389/fnhum.2017.00481 (2017). *
Cohen, M. X. _Analyzing Neural time Series Data: Theory and Practice_ (MIT Press, 2014). * Norcia, A. M., Appelbaum, L. G., Ales, J. M., Cottereau, B. R. & Rossion, B. The steady-state
visual evoked potential in vision research: A review. _J. Vis._ 15 (6), 4. https://doi.org/10.1167/15.6.4 (2015). Article PubMed PubMed Central Google Scholar * Lehmann, A., Arias, D. J.
& Schönwiesner, M. Tracing the neural basis of auditory entrainment. _Neuroscience_ 337, 306–314. https://doi.org/10.1016/j.neuroscience.2016.09.011 (2016). Article CAS PubMed Google
Scholar * Orozco Perez, H. D., Dumas, G. & Lehmann, A. Binaural beats through the auditory pathway: From brainstem to connectivity patterns. _Eneuro_ 7 (2), ENEURO0232–192020.
https://doi.org/10.1523/ENEURO.0232-19.2020 (2020). Article Google Scholar * Becher, A. K. et al. Intracranial electroencephalography power and phase synchronization changes during
monaural and binaural beat stimulation. _Eur. J. Neurosci._ 41 (2), 254–263. https://doi.org/10.1111/ejn.12760 (2015). Article PubMed Google Scholar * Tierney, A. & Kraus, N. Neural
entrainment to the rhythmic structure of music. _J. Cogn. Neurosci._ 27 (2), 400–408. https://doi.org/10.1162/jocn_a_00704 (2015). Article PubMed Google Scholar * Doelling, K. B. &
Poeppel, D. Cortical entrainment to music and its modulation by expertise. _Proc. Natl. Acad. Sci. U.S.A._ 112 (45), E6233–6242. https://doi.org/10.1073/pnas.1508431112 (2015). Article ADS
CAS PubMed PubMed Central Google Scholar * Nozaradan, S. Exploring how musical rhythm entrains brain activity with electroencephalogram frequency-tagging. _Philos. Trans. R. Soc. Lond.
B Biol. Sci._ 369 (1658), 20130393. https://doi.org/10.1098/rstb.2013.0393 (2014). Article PubMed PubMed Central Google Scholar * Collins, T., Laney, R., Willis, A. & Garthwaite, P.
H. Chopin, mazurkas and Markov: Making music in style with statistics. _Significance_ 8 (4), 154–159. https://doi.org/10.1111/j.1740-9713.2011.00519.x (2011). Article Google Scholar *
McConville, B. Noteflight as a web 2.0 tool for music theory pedagogy. _J. Music Theory Pedagogy_. 26 (1), 8 (2012). Article Google Scholar * London, J. _Hearing in Time: Psychological
Aspects of Musical Meter_ (Oxford University Press, 2012). * Bakker, I., van der Voordt, T., Vink, P. & de Boon, J. Pleasure, arousal, dominance: Mehrabian and Russell revisited. _Curr.
Psychol._ 33 (3), 405–421. https://doi.org/10.1007/s12144-014-9219-4 (2014). Article Google Scholar * Bradley, M. M. & Lang, P. J. Measuring emotion: The self-assessment manikin and
the semantic differential. _J. Behav. Ther. Exp. Psychiatry_. 25 (1), 49–59. https://doi.org/10.1016/0005-7916(94)90063-9 (1994). Article CAS PubMed Google Scholar * Wollman, I., Arias,
P., Aucouturier, J. J. & Morillon, B. Neural entrainment to music is sensitive to melodic spectral complexity. _J. Neurophysiol._ 123 (3), 1063–1071.
https://doi.org/10.1152/jn.00758.2018 (2020). Article PubMed PubMed Central Google Scholar * Nozaradan, S., Peretz, I. & Mouraux, A. Selective neuronal entrainment to the beat and
meter embedded in a musical rhythm. (2012). * Trost, W., Labbé, C. & Grandjean, D. Rhythmic entrainment as a musical affect induction mechanism. _Neuropsychologia_ 96, 96–110.
https://doi.org/10.1016/j.neuropsychologia.2017.01.004 (2017). Article Google Scholar * Fujioka, T., Trainor, L. J., Large, E. W. & Ross, B. Beta and gamma rhythms in human auditory
cortex during musical beat processing. _Ann. N. Y. Acad. Sci._ 1169 (1), 89–92. https://doi.org/10.1111/j.1749-6632.2009.04779.x (2009). Article ADS PubMed Google Scholar * Baltzell, L.
S., Srinivasan, R. & Richards, V. Hierarchical organization of melodic sequences is encoded by cortical entrainment. _NeuroImage_ 200, 490–500.
https://doi.org/10.1016/j.neuroimage.2019.06.054 (2019). Article PubMed Google Scholar * Marek, S. & Dosenbach, N. U. The frontoparietal network: Function, electrophysiology, and
importance of individual precision mapping. _Dialog. Clin. Neurosci._ 20 (2), 133–140 (2018). Article Google Scholar * Vuilleumier, P. & Trost, W. Music and emotions: From enchantment
to entrainment. _Ann. N. Y. Acad. Sci._ 1337 (1), 212–222. https://doi.org/10.1111/nyas.12676 (2015). Article ADS PubMed Google Scholar * Krause, A. E. & North, A. C. Pleasure,
arousal, dominance, and judgments about music in everyday life. _Psychol. Music_. 45 (3), 355–374. https://doi.org/10.1177/0305735616664214 (2017). Article Google Scholar * Helfrich, R.
F., Breska, A. & Knight, R. T. Neural entrainment and network resonance in support of top-down guided attention. _Curr. Opin. Psychol._ 29, 82–89.
https://doi.org/10.1016/j.copsyc.2018.12.016 (2019). Article PubMed PubMed Central Google Scholar * Hutchison, I. C. & Rathore, S. The role of REM sleep theta activity in emotional
memory. _Front. Psychol._ 6, 1439 (2015). Article PubMed PubMed Central Google Scholar * Hughes, S. W. & Crunelli, V. Thalamic mechanisms of EEG alpha rhythms and their pathological
implications. _Neurosci. Rev. J. Bringing Neurobiol. Neurol. Psychiatry_. 11 (4), 357–372. https://doi.org/10.1177/1073858405277450 (2005). Article Google Scholar * Mallik, A. & Russo,
F. A. The effects of music & auditory beat stimulation on anxiety: A randomized clinical trial. _PLoS ONE_. 17 (3), e0259312. https://doi.org/10.1371/journal.pone.0259312 (2022).
Article CAS PubMed PubMed Central Google Scholar * Padmanabhan, R., Hildreth, A. J. & Laws, D. A prospective, randomised, controlled study examining binaural beat audio and
pre-operative anxiety in patients undergoing general anaesthesia for day case surgery. _Anaesthesia_ 60 (9), 874–877. https://doi.org/10.1111/j.1365-2044.2005.04287.x (2005). Article CAS
PubMed Google Scholar * Wahbeh, H., Calabrese, C., Zwickey, H. & Zajdel, D. Binaural beat technology in humans: A pilot study to assess neuropsychologic, physiologic, and
electroencephalographic effects. _J. Altern. Complement. Med. (New York N Y)_. 13, 199–206. https://doi.org/10.1089/acm.2006.6201 (2007). Article Google Scholar * Weiland, T. J. et al.
Original sound compositions reduce anxiety in emergency department patients: A randomised controlled trial. _Med. J. Aust._ 195 (11–12), 694–698. https://doi.org/10.5694/mja10.10662 (2011).
Article PubMed Google Scholar * López-Caballero, F. & Escera, C. Binaural beat: A failure to enhance EEG power and emotional arousal. _Front. Hum. Neurosci._
https://doi.org/10.3389/fnhum.2017.00557 (2017). 11. Article PubMed PubMed Central Google Scholar * Critchley, H. D. & Garfinkel, S. N. Interoception and emotion. _Curr. Opin.
Psychol._ 17, 7–14 (2017). Article PubMed Google Scholar * Chen, J. L., Penhune, V. B. & Zatorre, R. J. Listening to musical rhythms recruits motor regions of the brain. _Cereb.
Cortex_. 18 (12), 2844–2854 (2008). Article PubMed Google Scholar * Witek, M. A. Rhythmic entrainment and embodied cognition. _Science-Music Borderlands_, 161–182. (2023). Download
references ACKNOWLEDGEMENTS We thank Hongjian Huang, Yufeng Xia who helped with data collection, and Dr. Yifan Xu for his expertise in music-related professional knowledge. FUNDING The work
described in this study was supported by the National Natural Science Foundation of China (62101324 and 32100885), the Fundamental Research Funds for the Central Universities (YG2025QNB13),
Shanghai Municipal Health Commission (2024ZZ2066), Shanghai Municipal Education Commission (2024AIYB012), Shanghai Mental Health Center (2020-QH-01, 2020-YJ01) and the Shanghai Sailing
Program (20YF1442000). AUTHOR INFORMATION Author notes * Jiaxin Ding and Xiaochen Zhang have contributed equally to this work. AUTHORS AND AFFILIATIONS * Shanghai Mental Health Center,
Shanghai Jiao Tong University, School of Medicine, Shanghai, China Jiaxin Ding, Xiaochen Zhang, Zhishan Hu, Yingying Tang & Yue Ding * Department of Psychosomatic Medicine, Shanghai East
Hospital Tongji University School of Medicine, Shanghai, China Jingjing Liu * Beijing Key Laboratory of Mental Disorders, National Clinical Research Center for Mental Disorders &
National Center for Mental Disorders, Beijing Anding Hospital, Capital Medical University, Beijing, China Zhi Yang * Mental Health Branch, China Hospital Development Institute, Shanghai Jiao
Tong University, Shanghai, China Yingying Tang & Yue Ding * Shanghai Key Laboratory of Psychotic Disorders, Shanghai, China Yingying Tang & Yue Ding Authors * Jiaxin Ding View
author publications You can also search for this author inPubMed Google Scholar * Xiaochen Zhang View author publications You can also search for this author inPubMed Google Scholar *
Jingjing Liu View author publications You can also search for this author inPubMed Google Scholar * Zhishan Hu View author publications You can also search for this author inPubMed Google
Scholar * Zhi Yang View author publications You can also search for this author inPubMed Google Scholar * Yingying Tang View author publications You can also search for this author inPubMed
Google Scholar * Yue Ding View author publications You can also search for this author inPubMed Google Scholar CONTRIBUTIONS Y.D., Z.Y. and YY. T. conceptualized the study. Y.D. also handled
data curation, formal analysis, methodology, visualization, and was responsible for writing the original draft. JX.D. contributed to formal analysis and writing the original draft. XC.Z.
secured funding and contributed to reviewing and editing the manuscript. JJ.L. was responsible for data collection, and ZS.H. provided resources. YY.T. supervised the study. Y.D. is the
guarantor. All authors reviewed and approved the final manuscript. CORRESPONDING AUTHORS Correspondence to Zhi Yang, Yingying Tang or Yue Ding. ETHICS DECLARATIONS CONSENT FOR PUBLICATION
All the participants gave informed consent to participate in the study before taking part. COMPETING INTERESTS The authors declare no competing interests. ETHICAL APPROVAL This study
involves human participants and was approved by the Shanghai Mental Health Center Institutional Review Board (SMHC-IRB ethics approval ID: 2021-45). Participants gave informed consent to
participate in the study before taking part. ADDITIONAL INFORMATION PUBLISHER’S NOTE Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional
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tonal sequences on neural oscillations and the impact on subjective emotion. _Sci Rep_ 15, 17462 (2025). https://doi.org/10.1038/s41598-025-98548-1 Download citation * Received: 19 October
2024 * Accepted: 14 April 2025 * Published: 20 May 2025 * DOI: https://doi.org/10.1038/s41598-025-98548-1 SHARE THIS ARTICLE Anyone you share the following link with will be able to read
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KEYWORDS * Rhythmic tonal sequence * Entrainment * Neural osillation * Subjective emotion