Neural Entrainment to Musical Pulse in Naturalistic Music Is Preserved in Aging: Implications for Music-Based Interventions.

Neural entrainment to musical rhythm is thought to underlie the perception and production of music. In aging populations, the strength of neural entrainment to rhythm has been found to be attenuated, particularly during attentive listening to auditory streams. However, previous studies on neural entrainment to rhythm and aging have often employed artificial auditory rhythms or limited pieces of recorded, naturalistic music, failing to account for the diversity of rhythmic structures found in natural music. As part of larger project assessing a novel music-based intervention for healthy aging, we investigated neural entrainment to musical rhythms in the electroencephalogram (EEG) while participants listened to self-selected musical recordings across a sample of younger and older adults. We specifically measured neural entrainment to the level of musical pulse-quantified here as the phase-locking value (PLV)-after normalizing the PLVs to each musical recording's detected pulse frequency. As predicted, we observed strong neural phase-locking to musical pulse, and to the sub-harmonic and harmonic levels of musical meter. Overall, PLVs were not significantly different between older and younger adults. This preserved neural entrainment to musical pulse and rhythm could support the design of music-based interventions that aim to modulate endogenous brain activity via self-selected music for healthy cognitive aging.

A Dynamical, Radically Embodied, and Ecological Theory of Rhythm Development.

Musical rhythm abilities-the perception of and coordinated action to the rhythmic structure of music-undergo remarkable change over human development. In the current paper, we introduce a theoretical framework for modeling the development of musical rhythm. The framework, based on Neural Resonance Theory (NRT), explains rhythm development in terms of resonance and attunement, which are formalized using a general theory that includes non-linear resonance and Hebbian plasticity. First, we review the developmental literature on musical rhythm, highlighting several developmental processes related to rhythm perception and action. Next, we offer an exposition of Neural Resonance Theory and argue that elements of the theory are consistent with dynamical, radically embodied (i.e., non-representational) and ecological approaches to cognition and development. We then discuss how dynamical models, implemented as self-organizing networks of neural oscillations with Hebbian plasticity, predict key features of music development. We conclude by illustrating how the notions of dynamical embodiment, resonance, and attunement provide a conceptual language for characterizing musical rhythm development, and, when formalized in physiologically informed dynamical models, provide a theoretical framework for generating testable empirical predictions about musical rhythm development, such as the kinds of native and non-native rhythmic structures infants and children can learn, steady-state evoked potentials to native and non-native musical rhythms, and the effects of short-term (e.g., infant bouncing, infant music classes), long-term (e.g., perceptual narrowing to musical rhythm), and very-long term (e.g., music enculturation, musical training) learning on music perception-action.

Integrating music-based interventions with Gamma-frequency stimulation: Implications for healthy ageing.

In recent years, music-based interventions (MBIs) have risen in popularity as a non-invasive, sustainable form of care for treating dementia-related disorders, such as Mild Cognitive Impairment (MCI) and Alzheimer's disease (AD). Despite their clinical potential, evidence regarding the efficacy of MBIs on patient outcomes is mixed. Recently, a line of related research has begun to investigate the clinical impact of non-invasive Gamma-frequency (e.g., 40 Hz) sensory stimulation on dementia. Current work, using non-human-animal models of AD, suggests that non-invasive Gamma-frequency stimulation can remediate multiple pathophysiologies of dementia at the molecular, cellular and neural-systems scales, and, importantly, improve cognitive functioning. These findings suggest that the efficacy of MBIs could, in theory, be enhanced by incorporating Gamma-frequency stimulation into current MBI protocols. In the current review, we propose a novel clinical framework for non-invasively treating dementia-related disorders that combines previous MBIs with current approaches employing Gamma-frequency sensory stimulation. We theorize that combining MBIs with Gamma-frequency stimulation could increase the therapeutic power of MBIs by simultaneously targeting multiple biomarkers of dementia, restoring neural activity that underlies learning and memory (e.g., Gamma-frequency neural activity, Theta-Gamma coupling), and actively engaging auditory and reward networks in the brain to promote behavioural change.

Beyond "consistent with" adaptation: Is there a robust test for music adaptation?

In their article, Mehr et al. conclude that the design features of music are consistent with adaptations for credible signaling. Although appealing to design may seem like a plausible basis for identifying adaptations, probing adaptive theories of music must be done at the genomic level and will require a functional understanding of the genomic, phenotypic, and fitness properties of music.

Bouncing the network: A dynamical systems model of auditory-vestibular interactions underlying infants' perception of musical rhythm.

Previous work suggests that auditory-vestibular interactions, which emerge during bodily movement to music, can influence the perception of musical rhythm. In a seminal study on the ontogeny of musical rhythm, Phillips-Silver and Trainor (2005) found that bouncing infants to an unaccented rhythm influenced infants' perceptual preferences for accented rhythms that matched the rate of bouncing. In the current study, we ask whether nascent, diffuse coupling between auditory and motor systems is sufficient to bootstrap short-term Hebbian plasticity in the auditory system and explain infants' preferences for accented rhythms thought to arise from auditory-vestibular interactions. First, we specify a nonlinear, dynamical system in which two oscillatory neural networks, representing developmentally nascent auditory and motor systems, interact through weak, non-specific coupling. The auditory network was equipped with short-term Hebbian plasticity, allowing the auditory network to tune its intrinsic resonant properties. Next, we simulate the effect of vestibular input (e.g., infant bouncing) on infants' perceptual preferences for accented rhythms. We found that simultaneous auditory-vestibular training shaped the model's response to musical rhythm, enhancing vestibular-related frequencies in auditory-network activity. Moreover, simultaneous auditory-vestibular training, relative to auditory- or vestibular-only training, facilitated short-term auditory plasticity in the model, producing stronger oscillator connections in the auditory network. Finally, when tested on a musical rhythm, models which received simultaneous auditory-vestibular training, but not models that received auditory- or vestibular-only training, resonated strongly at frequencies related to their "bouncing," a finding qualitatively similar to infants' preferences for accented rhythms that matched the rate of infant bouncing.

Modeling infants' perceptual narrowing to musical rhythms: neural oscillation and Hebbian plasticity.

Previous research suggests that infants' perception of musical rhythm is fine-tuned to culture-specific rhythmic structures over the first postnatal year of human life. To date, however, little is known about the neurobiological principles that may underlie this process. In the current study, we used a dynamical systems model featuring neural oscillation and Hebbian plasticity to simulate infants' perceptual learning of culture-specific musical rhythms. First, we demonstrate that oscillatory activity in an untrained network reflects the rhythmic structure of either a Western or a Balkan training rhythm in a veridical fashion. Next, during a period of unsupervised learning, we show that the network learns the rhythmic structure of either a Western or a Balkan training rhythm through the self-organization of network connections. Finally, we demonstrate that the learned connections affect the networks' response to violations to the metrical structure of native and nonnative rhythms, a pattern of findings that mirrors the behavioral data on infants' perceptual narrowing to musical rhythms.

Musical Experience, Sensorineural Auditory Processing, and Reading Subskills in Adults.

Developmental research suggests that sensorineural auditory processing, reading subskills (e.g., phonological awareness and rapid naming), and musical experience are related during early periods of reading development. Interestingly, recent work suggests that these relations may extend into adulthood, with indices of sensorineural auditory processing relating to global reading ability. However, it is largely unknown whether sensorineural auditory processing relates to specific reading subskills, such as phonological awareness and rapid naming, as well as musical experience in mature readers. To address this question, we recorded electrophysiological responses to a repeating click (auditory stimulus) in a sample of adult readers. We then investigated relations between electrophysiological responses to sound, reading subskills, and musical experience in this same set of adult readers. Analyses suggest that sensorineural auditory processing, reading subskills, and musical experience are related in adulthood, with faster neural conduction times and greater musical experience associated with stronger rapid-naming skills. These results are similar to the developmental findings that suggest reading subskills are related to sensorineural auditory processing and musical experience in children.

Frequency-dependent fine structure in the frequency-following response: The byproduct of multiple generators.

The frequency-following response (FFR) is an auditory-evoked response recorded at the scalp that captures the spectrotemporal properties of tonal stimuli. Previous investigations report that the amplitude of the FFR fluctuates as a function of stimulus frequency, a phenomenon thought to reflect multiple neural generators phase-locking to the stimulus with different response latencies. When phase-locked responses are offset by different latencies, constructive and destructive phase interferences emerge in the volume-conducted signals, culminating in an attenuation or amplification of the scalp-recorded response in a frequency-specific manner. Borrowing from the literature on the audiogram and otoacoustic emissions (OAEs), we refer to this frequency-specific waxing and waning of the FFR amplitude as fine structure. While prior work on the human FFR was limited by small sets of stimulus frequencies, here, we provide the first systematic investigation of FFR fine structure using a broad stimulus set (90 + frequencies) that spanned the limits of human pitch perception. Consistent with predictions, the magnitude of the FFR response varied systematically as a function of stimulus frequency between 16.35 and 880 Hz. In our dataset, FFR high points (local maxima) emerged at ∼44, 87, 208, and 415 Hz with FFR valleys (local minima) emerging ∼62, 110, 311, and 448 Hz. To investigate whether these amplitude fluctuations are the result of multiple neural generators with distinct latencies, we created a theoretical model of the FFR that included six putative generators. Based on the extant literature on the sources of the FFR, our model adopted latencies characteristic of the cochlear microphonic (0 ms), cochlear nucleus (∼1.25 ms), superior olive (∼3.7 ms), and inferior colliculus (∼5 ms). In addition, we included two longer latency putative generators (∼13 ms, and ∼25 ms) reflective of the characteristic latencies of primary and non-primary auditory cortical structures. Our model revealed that the FFR fine structure observed between 16.35 and 880 Hz can be explained by the phase-interaction patterns created by six generators with relative latencies spaced between 0 and 25 ms. In addition, our model provides confirmatory evidence that both subcortical and cortical structures are activated by low-frequency (<100 Hz) tones, with the cortex being less sensitive to frequencies > 100 Hz. Collectively, these findings highlight (1) that the FFR is a composite response; (2) that the FFR at any given frequency can reflect activity from multiple generators; (3) that the fine-structure pattern between 16.35 and 880 Hz is the collective outcome of short- and long-latency generators; (4) that FFR fine structure is epiphenomenal in that it reflects how volume-conducted electrical potentials originating from different sources with different latencies interact at scalp locations, not how these different sources actually interact in the brain; and (5) that as a byproduct of these phase-interaction patterns low-amplitude responses will emerge at some frequencies, even when the underlying generators are fully functioning. We believe these findings call for a re-examination of how FFR amplitude is interpreted in both clinical and experimental contexts.

Linking prenatal experience to the emerging musical mind.

The musical brain is built over time through experience with a multitude of sounds in the auditory environment. However, learning the melodies, timbres, and rhythms unique to the music and language of one's culture begins already within the mother's womb during the third trimester of human development. We review evidence that the intrauterine auditory environment plays a key role in shaping later auditory development and musical preferences. We describe evidence that externally and internally generated sounds influence the developing fetus, and argue that such prenatal auditory experience may set the trajectory for the development of the musical mind.

Effects of perceptual experience on children's and adults' perception of unfamiliar rhythms.

Rhythm and meter are fundamental components of music that are universal yet also culture specific. Although simple, isochronous meters are preferred and more readily discriminated than highly complex, nonisochronous meters, moderately complex nonisochronous meters do not pose a problem for listeners who are exposed to them from a young age. The present work uses a behavioral task to examine the ease with which listeners of various ages acquire knowledge of unfamiliar metrical structures from passive exposure. We examined perception of familiar (Western) rhythms with an isochronous meter and unfamiliar (Balkan) rhythms with a nonisochronous meter. We compared discrimination by American children (5 to 11 years) and adults before and after a 2-week period of at-home listening to nonisochronous meter music from Bulgaria. During the first session, listeners of all ages exhibited superior discrimination of isochronous than in nonisochronous melodies. Across sessions, this asymmetry declined for young children but not for older children and adults.