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Brainwaves: Scientific Exploration and Tuning Fork Applications

  • Writer: Tuning Chi
    Tuning Chi
  • Aug 10
  • 14 min read

Brainwaves are all the rage — increasingly mentioned and often credited with a wide range of benefits.I wanted to cut through the common misconceptions and explore what science actually has to say about them.Once confined to neurophysiology laboratories, their study has seen a resurgence in recent years, driven by research into their cognitive, emotional, and therapeutic applications.This article offers a synthesis of current knowledge, based on recent scientific publications, and explores how therapeutic tuning forks can provide a tangible way to interact with these fundamental rhythms.


1. Understanding Brainwaves and Their Synchronization


1.1 What is a Brainwave?

Brainwaves are electrical oscillations produced by the synchronized activity of groups of neurons. They are classified according to their frequency:

  • Delta (0.5–4 Hz) – Deep sleep, regeneration.

  • Theta (4–8 Hz) – Memory processing, daydreaming, relaxation.

  • Alpha (8–12 Hz) – Calm alertness, reduced distractions.

  • Beta (12–30 Hz) – Active attention, conscious thought.

  • Gamma (>30 Hz, often around 40 Hz) – Coordination between different brain regions, working memory, advanced cognition.

These brainwave types coexist at all times, but their relative dominance varies depending on states of wakefulness, focus, or relaxation.


1.2 Synchronizing Brainwaves Through Rhythmic Stimulation

The brain has a natural ability to synchronize its brainwaves with external frequencies when they are delivered in a regular, rhythmic manner. This phenomenon occurs through auditory stimulation (binaural beats), visual stimulation (light patterns), or mechanical stimulation (vibrations applied to the body).

In their study "An Integrative Review of Brainwave Entrainment Benefits for Human Health" (Cunha et al., 2025), the authors analyzed 84 scientific studies on the effects of brainwave synchronization through external stimulation. The results highlight benefits in:

  • Pain reduction

  • Improved sleep quality

  • Stress, anxiety, and depression management

  • Enhancement of cognitive functions (focus, memory)

  • Support for patients with neurodegenerative disorders

These effects vary depending on the targeted frequencies (Delta, Theta, Alpha, Gamma), as each brainwave band is associated with specific physiological functions.


1.3 Brainwaves and Anesthesia: Reducing Sedative Requirements

Schmid et al. (2020) evaluated the impact of rhythmic stimulation (binaural beats and light stimuli) on propofol requirements during pediatric surgeries.

Protocol:

  • 49 children divided into two groups: one receiving stimulation, the other not.

  • Targeted frequencies were those associated with relaxation and deep sleep (theta and delta).

Results:

  • The stimulated group required significantly lower doses of propofol (3.0 mg/kg/h vs. 4.2 mg/kg/h).

  • Sedation depth remained equivalent between groups.

These findings suggest that synchronizing brainwaves with slower frequencies can promote a sedative state, thereby reducing the amount of medication needed.


1.4 Brainwaves and Tinnitus: Reducing Emotional Handicap

Prakash & Konadath (2025) studied the effect of binaural beats on patients with tinnitus who had normal hearing. Participants were exposed to different frequencies (delta, theta, alpha) and compared to a standard sound-masking intervention (white noise).

After three months of daily stimulation:

  • Groups receiving binaural beats showed a greater reduction in tinnitus-related handicap.

  • They also reported decreased stress and depression levels.

  • Perceived quality of life improved.

The authors suggest that synchronizing brainwaves at specific frequencies may help rebalance neural circuits involved in sound perception and emotional regulation.


1.5 Brainwaves and Cognitive Disorders: Specific Alterations by PathologyBaşar & Güntekin (2013) analyzed alterations in brainwaves across several neuropsychiatric disorders:

  • Schizophrenia: reduced gamma responses at 40 Hz, desynchronization in delta, theta, and alpha bands, impacting memory and attention.

  • Alzheimer’s disease & MCI: disruption of theta and alpha bands, loss of gamma synchronization.

  • ADHD: deficits in theta and alpha during sustained attention tasks.

These findings show that brainwaves can serve as dynamic biomarkers to refine the diagnosis of cognitive pathologies. Modulating these rhythms opens perspectives for targeted therapeutic strategies.


1.6 Brainwaves and Sleep: Optimizing Mental Recovery

Abeln et al. (2014) conducted a study on young elite soccer players to evaluate the impact of auditory stimulation (binaural beats between 2–8 Hz) during sleep.

Protocol:

  • 15 players were exposed each night for 8 weeks to Delta/Theta binaural beats.

  • 15 sports students served as a control group (no stimulation).

  • Weekly questionnaires assessed subjective sleep quality and morning motivation levels.

Results:

  • The stimulated group showed a significant improvement in perceived sleep quality.

  • Increased feelings of mental rest and higher motivation upon waking were observed.

  • No immediate effect on perceived muscle fatigue, though the authors note that physical benefits might appear over a longer term.

This study highlights the potential of rhythmic stimulation to optimize mental recovery, particularly in contexts requiring high cognitive performance.


2. Binaural Beats

Binaural beats are perceived when listening to two slightly different frequencies in each ear via headphones. The brain then perceives a third frequency corresponding to the difference between the two.


2.1 Comparison with Monaural Beats

In contrast, monaural beats result from the physical superposition of two frequencies in the air or within the same auditory channel.

Binaural Beat

Monaural Beat

Two distinct frequencies, one in each ear

Two frequencies mixed into the same signal

Neuro-acoustic phenomenon (central perception)

Acoustic phenomenon (directly perceived in the ear)

Requires stereo headphones

Works with headphones or loudspeakers


In summary, binaural beats seem to induce specific neurofunctional effects through central auditory integration, whereas monaural beats act mainly through direct modulation of the acoustic signal.


2.2 Effects of Binaural Beats on Brain Activity

Several studies have explored the underlying brain mechanisms of binaural beats:

  • Ross et al. (2014) – EEG study comparing binaural and monaural beats. The authors observed that BB activated specific central auditory processing mechanisms, particularly in the temporal and parietal regions, with dynamics distinct from those of monaural beats.

  • Solcà et al. (2015) – EEG study on participants exposed to alpha-band BB (10 Hz). Result: increased interhemispheric coherence in this frequency band, suggesting improved functional connectivity between the auditory cortices.

  • Scala et al. (2025) – Functional MRI comparison between BB and monaural beats at 6 Hz. BB produced specific activation of the cuneus and precuneus, as well as mild modulation of autonomic nervous system parameters (heart rate variability). Monaural beats did not show these targeted effects.

These findings highlight that binaural beats can influence brain activity and functional connectivity in ways not observed with monaural beats.


3. Brainwaves: Observed Effects by Targeted Band

Each brainwave band is associated with distinct physiological and psychological states. The following sections summarize scientific findings for each of the main frequency ranges, along with potential therapeutic applications.


3.1 — Alpha Waves (8–12 Hz)

Alpha waves are associated with a calm yet alert state, reduced anxiety, facilitation of learning, and decreased pain perception.They are among the most studied brainwave frequencies in the context of non-invasive modulation of brain activity, particularly through sound stimulation. Below are three scientific studies that have evaluated their impact in different clinical and cognitive contexts.


a) Alpha and Pain Reduction

Ajo et al. (2019) investigated the effect of alpha binaural beats (10 Hz) on the perception of pain induced by a thermal stimulus.

  • Method: 29 participants were exposed to a moderately painful thermal stimulus while listening either to musical audio containing 10 Hz BB or to the same musical audio alone (control).

  • Results: The BB group showed a significant increase in pain tolerance threshold (+8% on average), as well as a reduction in perceived intensity.

  • Observation: The authors suggest that the analgesic effect may be related to increased interhemispheric coherence in the alpha band, facilitating emotional regulation and central pain modulation.


b) Alpha and Preoperative Anxiety

Padmanabhan et al. (2005) evaluated the effect of alpha BB (10 Hz) on preoperative anxiety in hospitalized patients prior to surgery.

  • Method: 108 patients were divided into 3 groups: alpha BB + relaxing music, music only, or no stimulation.

  • Results: Significant reduction in STAI (State-Trait Anxiety Inventory) scores in the BB group (−26%), greater than in the music-only group (−11%), with no notable change in the control group.

  • Observation: The authors noted that the anxiolytic effect was observable after only 30 minutes of exposure, suggesting that this could be a simple complementary tool in preoperative preparation protocols.


c) Alpha and Learning / Memory

Kennerly (1994) tested the effect of alpha BB on the learning of word lists.

  • Method: 30 participants listened either to 10 Hz BB or to pink noise while performing a verbal learning task.

  • Results: Improvement of 13% in both immediate and delayed recall in the BB group compared to the control group.

  • Observation: This improvement may be linked to the calm alertness induced by alpha waves, optimizing memory consolidation.


3.2 — Theta Waves (4–8 Hz)

Theta waves are linked to deep relaxation, daydreaming, memory access, and creativity. They also play a role in certain cognitive processes such as memory consolidation — the process by which recently acquired memories are stabilized and stored in long-term memory — and emotional regulation.Below are three scientific studies that have explored the impact of theta waves in various contexts, ranging from brain connectivity functions to decision-making and pain processing.


a) Theta and Brain Connectivity in Meditation

Aftanas & Golocheikine (2001) studied theta wave activity recorded in the frontal lobe (the anterior part of the skull, above the medial prefrontal cortex) and functional connectivity during “open monitoring” meditation in experienced practitioners and novices.

  • Method: EEG recordings on 16 experienced meditators and 16 novices during standardized meditation sessions. Analysis of power and connectivity changes.

  • Results: Meditators showed a significant increase in frontal-midline theta power (+21% compared to novices, p < 0.05) and higher fronto-temporal connectivity (+18%, p < 0.05).

  • Observation: Experienced meditators displayed greater coherence between EEG signals in the frontal and parietal regions, indicating more efficient communication between these areas. The authors suggest that enhanced theta connectivity may be a neurophysiological marker of the emotional stability observed in experienced practitioners.


b) Theta and Decision-Making in Parkinson’s Patients

Singh et al. (2021) explored theta activity measured in the frontal-midline region (central anterior head area) during decision-making tasks in patients with Parkinson’s disease.

  • Method: EEG recordings from 25 Parkinson’s patients and 25 healthy controls during an Iowa Gambling Task.

  • Results: Significant reduction in frontal-midline theta power in Parkinson’s patients (−15%, p < 0.05) compared to controls. This decrease was correlated with poorer decision-making performance (r = 0.46, p < 0.05).

  • Observation: The authors suggest that enhancing frontal-midline theta activity, particularly via sound stimulation or neurofeedback, could help improve certain executive functions impaired in Parkinson’s disease.


c) Theta and Pain Modulation

Barratt et al. (2011) examined the effects of auditory stimulation with theta BB (6 Hz) on experimentally induced pain perception.

  • Method: 24 participants were exposed to a moderately painful thermal stimulus while listening either to musical audio containing 6 Hz BB or to the same musical audio alone (control). EEG was recorded during the experiment.

  • Results: Significant increase in pain tolerance (+12%, p < 0.05) and reduction in perceived intensity (−9%, p < 0.05) in the BB group compared to the control. EEG showed increased theta power in the frontal and temporal regions.

  • Observation: The authors propose that theta waves may promote a state of sensory dissociation or distraction, thereby reducing pain perception.


3.3 — Delta Waves (0.5–4 Hz)

Delta waves are the slowest in the brainwave spectrum. They dominate during slow-wave deep sleep (stage N3) and are associated with tissue regeneration, mental recovery, and reduced pain perception. Below are two studies that explored the effects of delta wave–enhancing stimulation in specific clinical contexts.


a) Delta and Tinnitus: Reducing Handicap and Anxiety

Prakash & Konadath (2024) evaluated the impact of auditory stimulation using binaural beats at delta frequency (4 Hz) in individuals with tinnitus and normal hearing.

Method: 50 participants were divided into an experimental group (4 Hz BB, 20 min/day for 30 days) and a control group (white noise, standard masking).

Results:

  • Average 24% reduction in tinnitus handicap score (THI – Tinnitus Handicap Inventory, p < 0.05).

  • Average 18% decrease in anxiety symptoms and 15% decrease in depressive symptoms (validated questionnaires, p < 0.05).

  • Greater improvement in the BB group compared to the white noise group.

Observation: The authors suggest that delta stimulation may help modulate neuronal and emotional activity involved in tinnitus perception, while noting that some participants showed only limited response.


b) Delta and Sleep: Improving Quality and Mood

Dabiri et al. (2022) conducted a pilot study on the effect of delta binaural beats (4 Hz) listened to for 90 minutes before or during sleep in 20 healthy students.

Method: Two-week protocol — one control week with no intervention and one experimental week with delta BB exposure. Sleep was tracked with a sleep diary, and mood was assessed with the Profile of Mood States (POMS) questionnaire.

Results:

  • Reduced number of nocturnal awakenings.

  • Increased total sleep duration.

  • Improved subjective sleep quality.

  • Enhanced mood upon waking, with reduced anxiety and anger (no significant difference for other emotional parameters).

Observation: The authors suggest that delta stimulation could be a simple, low-cost, side-effect-free intervention to improve sleep quality and support emotional regulation.


3.4 — Gamma Waves (30–100 Hz)

Gamma waves are the fastest in the brainwave spectrum. They are associated with higher cognitive functions such as focused attention, working memory, conscious perception, creativity, and efficient coordination between different brain regions. In neuroscience, they are also considered a marker of neural coherence and multisensory integration.

In recent years, interest in gamma waves has increased significantly, particularly for their potential neuroprotective role in conditions such as Alzheimer’s disease (AD) and post-stroke cognitive disorders. The following studies illustrate their cognitive, emotional, and physiological effects, as well as the neuronal and glial mechanisms involved.


a) Gamma and Cognitive Functions: Memory, Mood, Attention

Sharpe et al. (2020) assessed the impact of 40 Hz gamma binaural beats on memory, mood, and cognitive abilities in 9 participants divided into three groups (40 Hz, 25 Hz, 100 Hz).

Method: Eight sessions over four weeks; assessments before and after 5 minutes of binaural stimulation at the assigned frequency.

Results:

  • Average cognitive score improvement of +10% (75% → 85%) for the 40 Hz group (p = 0.076).

  • Significant improvement in memory scores by +8% (87% → 95%, p = 0.0027).

  • Improved mood (negative correlation between questionnaire scores and mood, indicating reduced anxiety and irritability).

Observation: The authors note that 40 Hz gamma stimulation may enhance neuronal coherence and optimise communication between brain regions involved in memory and emotional regulation.


b) Gamma and Neuroprotection: Alzheimer’s and Neurodegenerative Disorders

Adaikkan et al. (2019) demonstrated that 40 Hz sensory stimulation (GENUS: Gamma Entrainment Using Sensory stimuli) could reduce amyloid deposits and phosphorylated tau protein in several mouse models of neurodegeneration. Recordings showed that gamma stimulation engaged the visual cortex, hippocampus, and prefrontal cortex.

Method: Daily exposure to 40 Hz visual stimulation (GENUS) from early disease stages.

Results (after gamma stimulation):

  • Preservation of neuronal and synaptic density in multiple brain regions.

  • Improved cognitive performance (memory and spatial orientation tests).

  • Reduced microglial inflammation and decreased neuronal DNA damage.

Observation: The authors suggest that gamma stimulation activates neuroprotective pathways by modulating the activity of neurons and glial cells (astrocytes and microglia), thereby slowing the progression of neurodegenerative lesions.


c) Gamma and Cellular Modulation: Neurons, Astrocytes, and Microglia

Adaikkan & Tsai (2020) further explored the cellular mechanisms involved in gamma stimulation, studying the effect of 40 Hz entrainment on different cell types and cerebral vascularisation.

Method: 40 Hz sensory stimulation in animal models, with neurophysiological, histological, and molecular analyses targeting neurons, glial cells, and vascular parameters.

Results:

  • Coordinated activation of a neural circuitry network including excitatory neurons, inhibitory interneurons, and glial cells (astrocytes and microglia).

  • Microglia: shift from a pro-inflammatory (M1) to an anti-inflammatory (M2) phenotype, promoting clearance of neuronal debris.

  • Neurons: improved synaptic synchronisation and increased interregional neuronal coherence.

  • Astrocytes: enhanced metabolic support and modulation of active synapses.

  • Vascular effect: increased cerebral blood flow and reduced inflammation via nitric oxide release, improving neuronal oxygenation and nutrition.

Observation: The authors suggest that these combined neuroprotective, anti-inflammatory, and vascular effects could help slow the progression of neurodegenerative diseases such as Alzheimer’s.


d) Gamma and Post-Stroke Cognitive Disorders

Adaikkan & Tsai (2020) also examined gamma stimulation in the context of post-stroke cognitive deficits, analysing its impact on neurons, glial cells, cerebral vascularisation, and neuronal connectivity.

Method: 40 Hz sensory stimulation in animal models, with neurophysiological, histological, and molecular assessments.

Results:

  • Coordinated activation of neuronal and glial networks (astrocytes and microglia).

  • Shift of microglia to an anti-inflammatory profile, improved neuronal coherence, and enhanced synaptic plasticity.

  • Positive vascular effect with increased cerebral blood flow and better neuronal oxygenation.

  • Strengthened functional connectivity between frontal, parietal, and hippocampal regions.

Observation: The authors suggest that this combination of neuroprotective, anti-inflammatory, and vascular effects could slow the progression of neurodegenerative diseases.



4 — Tuning Forks to Activate and Synchronize Brainwaves

Tuning forks are ideal tools for targeting specific brainwave bands. Depending on the chosen configuration, they can act via auditory pathways or through direct mechanical conduction.

Several frequency combinations and application methods are presented here, covering both dedicated “brain tuner” forks and more commonly used therapeutic tuning forks.


4.1 – Choosing the Tuning Forks


A) “Brain Tuners” Series

Unweighted tuning forks, specifically tuned to generate a binaural beat that directly corresponds to a target brainwave band (delta, alpha, theta, gamma). They are used in pairs, with one fork placed near each ear so that the brain perceives a frequency difference matching the desired brainwave.


B) “Classic” Tuning Fork Combinations

Two more commonly used tuning forks (weighted or unweighted) are paired so that their frequency difference corresponds to the target brainwave. Here are some suggested combinations:

  • Alpha (8–12 Hz): 128 Hz (Pythagorean frequency / rounded Schumann resonance at 8 Hz) + 136.10 Hz (Earth year frequency) → 8.10 Hz, within the alpha range.


    Alternative : 125.28 Hz (exact 16th harmonic of Schumann resonance: 7.83 × 16) + 136.10 Hz → 10.82 Hz, also within alpha range and producing comparable effects.

  • Theta (4–8 Hz) : 64 Hz (octave below Schumann resonance) + 68.05 Hz (octave below Earth year frequency) → 4.05 Hz, mid-theta range.

  • Delta (<4 Hz) : 128 Hz (Pythagorean / rounded Schumann) + 125.28 Hz (Schumann × 16) → 2.72 Hz.

  • Gamma (~40 Hz) : 256 Hz (C) + 296 Hz (unweighted) → 40 Hz.


    Alternative with weighted forks: 64 Hz (Schumann) + 104 Hz → 40 Hz.


    Note : 296 Hz and 104 Hz forks are not commonly used in practice but are commercially available and proposed here as possible combinations.


Summary Table of Commonly Used Therapeutic Combinations for Targeting Brainwave Bands


Brainwave Band

Suggested Combinations

Frequency Difference

Alpha (8–12 Hz)

128 Hz (Pythagorean / rounded Schumann) + 136.10 Hz (Earth year) or 125.28 Hz (Schumann ×16) + 136.10 Hz (Earth year)

8.10 Hz 10.82 Hz

Theta (4–8 Hz)

64 Hz (octave below Schumann) + 68.05 Hz (octave below Earth year)

4.05 Hz

Delta (<4 Hz)

128 Hz (Pythagorean / rounded Schumann) + 125.28 Hz (Schumann ×16)

2.72 Hz

Gamma (~40 Hz)

256 Hz (C) + 296 Hz or 64 Hz (Schumann) + 104 Hz

40 Hz


4.2 – Two Application Methods


a) Near the Ears

One tuning fork is placed near each ear (without direct contact with the pinna). The difference between the two perceived frequencies creates a binaural beat via the auditory pathway, which induces synchronization to the target brainwave frequency.


b) On the Body

Weighted tuning forks can also be applied directly to the body. In this method, the tips of the two stems are brought into contact, forming a “V,” to transmit vibration mechanically through the tissues.

Unlike ear application, no binaural beat is generated. The intended effect relies on somatosensory stimulation: the vibration travels through tissues, activating deep receptors (fascia, bones, joints) and providing direct sensory feedback.

This physical interaction:

  • Reinforces sensorimotor alignment by combining tactile and auditory perception.

  • Encourages conscious engagement by making the frequency “palpable” in the body.

  • Simultaneously stimulates multiple sensory networks (auditory, tactile, proprioceptive), potentially enhancing the brain’s integration of the target frequency.


Conclusion

Recent research confirms that brainwave modulation is no longer just a laboratory curiosity but a genuine therapeutic field. Whether aiming to promote relaxation, support cognitive recovery, or accompany certain medical conditions, rhythmic stimulation — auditory, visual, or mechanical — now offers promising possibilities.

Tuning forks, thanks to their precision and ability to act both through auditory pathways and tissue conduction, provide a simple and tangible way to resonate with these fundamental rhythms.

The potential of brainwaves and their broad application range make them a valuable ally for enriching a therapeutic practice.



Références Bibliographiques

  1. Schmid, W. et al. (2020). Brainwave entrainment to minimise sedative drug doses in paediatric surgery: a randomised controlled trial. British Journal of Anaesthesia.

  2. Herweg, N. A., Solomon, E. A., Kahana, M. J. (2020). Theta Oscillations in Human Memory. Trends in Cognitive Sciences.

  3. Cole, R. C. et al. (2025). Theta-frequency subthalamic nucleus stimulation increases decision threshold. Brain Stimulation.

  4. Okabe, N. et al. (2025). Theta Frequency Electromagnetic Stimulation Enhances Functional Recovery After Stroke. Neurorehabilitation & Neural Repair.

  5. Prakash, P., Konadath, S. (2024). Outcome measures of brainwave entrainment using delta wave stimulation in individuals with tinnitus having normal hearing sensitivity. Audiology Research.

  6. Dabiri, R. et al. (2022). The effect of auditory stimulation using delta binaural beat for a better sleep and post-sleep mood: A pilot study. Noise & Health.

  7. Wiesman, A. I., Wilson, T. W. (2019). Alpha Frequency Entrainment Reduces the Effect of Visual Distractors. Journal of Cognitive Neuroscience.

  8. Halpin, S. J. et al. (2025). Pre-sleep alpha brain entrainment by audio or visual stimulation for chronic widespread pain and sleep disturbance: A randomised crossover feasibility trial. Pain Reports.

  9. Shehani, F. A. et al. (2024). Effectiveness of Preoperative Alpha Wave Entrainment in Pediatric Dental Patients: A Randomized Controlled Trial. Journal of Dental Research.

  10. Chen, X. et al. (2025). Unleashing the potential: 40 Hz multisensory stimulation therapy for cognitive impairment. Neuroscience Insights.

  11. Sharpe, R. L. S. et al. (2020). Gamma entrainment frequency affects mood, memory and cognition: an exploratory pilot study. Brain Sciences.

  12. Adaikkan, C. et al. (2019). Gamma Entrainment Binds Higher-Order Brain Regions and Offers Neuroprotection. Neuron.

  13. Adaikkan, C., Tsai, L.-H. (2020). Gamma Entrainment: Impact on Neurocircuits, Glia, and Therapeutic Opportunities. Trends in Neurosciences.

  14. Kim, H.-S. (2021). Can gamma entrainment of the brain rhythms prevent or alleviate Alzheimer’s disease? Frontiers in Neuroscience.

  15. Sahu, M. et al. (2024). Harnessing Brainwave Entrainment: A Non-invasive Strategy To Alleviate Neurological Disorder Symptoms. Brain and Behavior.

  16. Başar, E. (2013). A review of gamma oscillations in healthy subjects and in cognitive impairment. International Journal of Psychophysiology.

  17. Attokaren, M. K. et al. (2023). BrainWAVE: A Flexible Method for Noninvasive Stimulation of Brain Rhythms across Species. Scientific Reports.

  18. Engelbregt, H. et al. (2021). Effects of binaural and monaural beat stimulation on attention and EEG. Experimental Brain Research.

  19. Ross, B. et al. (2014). Human cortical responses to slow and fast binaural beats reveal multiple mechanisms of binaural hearing. PLOS ONE.

  20. Solcà, M. et al. (2015). Binaural beats increase interhemispheric alpha-band coherence between auditory cortices. Scientific Reports.

  21. Scala, I. et al. (2025). Analysis of the effect of 6-Hz binaural beats on electroencephalographic and autonomic parameters of healthy individuals: An exploratory study. Journal of Neural Transmission.




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