Takako Fujioka

Musical Training Enhances Automatic Encoding of Melodic Contour and Interval Structure

In music, melodic information is thought to be encoded in two forms, a contour code (up/down pattern of pitch changes) and an interval code (pitch distances between successive notes). A recent study recording the mismatch negativity (MMN) evoked by pitch contour and interval deviations in simple melodies demonstrated that people with no formal music education process both contour and interval information in the auditory cortex automatically. However, it is still unclear whether musical experience enhances both strategies of melodic encoding. We designed stimuli to examine contour and interval information separately. In the contour condition there were eight different standard melodies (presented on 80 of trials), each consisting of five notes all ascending in pitch, and the corresponding deviant melodies (20) were altered to descending on their final note. The interval condition used one five-note standard melody transposed to eight keys from trial to trial, and on deviant trials the last note was raised by one whole tone without changing the pitch contour. There was also a control condition, in which a standard tone (990.7 Hz) and a deviant tone (1111.0 Hz) were presented. The magnetic counterpart of the MMN (MMNm) from musicians and nonmusicians was obtained as the difference between the dipole moment in response to the standard and deviant trials recorded by magnetoencephalography. Significantly larger MMNm was present in musicians in both contour and interval conditions than in nonmusicians, whereas MMNm in the control condition was similar for both groups. The interval MMNm was larger than the contour MMNm in musicians. No hemispheric difference was found in either group. The results suggest that musical training enhances the ability to automatically register abstract changes in the relative pitch structure of melodies.

Internalized Timing of Isochronous Sounds Is Represented in Neuromagnetic Beta Oscillations

Moving in synchrony with an auditory rhythm requires predictive action based on neurodynamic representation of temporal information. Although it is known that a regular auditory rhythm can facilitate rhythmic movement, the neural mechanisms underlying this phenomenon remain poorly understood. In this experiment using human magnetoencephalography, 12 young healthy adults listened passively to an isochronous auditory rhythm without producing rhythmic movement. We hypothesized that the dynamics of neuromagnetic beta-band oscillations (∼20 Hz)—which are known to reflect changes in an active status of sensorimotor functions—would show modulations in both power and phase-coherence related to the rate of the auditory rhythm across both auditory and motor systems. Despite the absence of an intention to move, modulation of beta amplitude as well as changes in cortico-cortical coherence followed the tempo of sound stimulation in auditory cortices and motor-related areas including the sensorimotor cortex, inferior-frontal gyrus, supplementary motor area, and the cerebellum. The time course of beta decrease after stimulus onset was consistent regardless of the rate or regularity of the stimulus, but the time course of the following beta rebound depended on the stimulus rate only in the regular stimulus conditions such that the beta amplitude reached its maximum just before the occurrence of the next sound. Our results suggest that the time course of beta modulation provides a mechanism for maintaining predictive timing, that beta oscillations reflect functional coordination between auditory and motor systems, and that coherence in beta oscillations dynamically configure the sensorimotor networks for auditory-motor coupling.

Beta and gamma rhythms in human auditory cortex during musical beat processing.

We examined β‐ (∼20 Hz) and γ‐ (∼40 Hz) band activity in auditory cortices by means of magnetoencephalography (MEG) during passive listening to a regular musical beat with occasional omission of single tones. The β activity decreased after each tone, followed by an increase, thus forming a periodic modulation synchronized with the stimulus. The β decrease was absent after omissions. In contrast, γ‐band activity showed a peak after tone and omission, suggesting underlying endogenous anticipatory processes. We propose that auditory β and γ oscillations have different roles in musical beat encoding and auditory–motor interaction.

Automatic Encoding of Polyphonic Melodies in Musicians and Nonmusicians

In music, multiple musical objects often overlap in time. Western polyphonic music contains multiple simultaneous melodic lines (referred to as voices) of equal importance. Previous electrophysiological studies have shown that pitch changes in a single melody are automatically encoded in memory traces, as indexed by mismatch negativity (MMN) and its magnetic counterpart (MMNm), and that this encoding process is enhanced by musical experience. In the present study, we examined whether two simultaneous melodies in polyphonic music are represented as separate entities in the auditory memory trace. Musicians and untrained controls were tested in both magnetoencephalogram and behavioral sessions. Polyphonic stimuli were created by combining two melodies (A and B), each consisting of the same five notes but in a different order. Melody A was in the high voice and Melody B in the low voice in one condition, and this was reversed in the other condition. On 50 of trials, a deviant final (5th) note was played either in the high or in the low voice, and it either went outside the key of the melody or remained within the key. These four deviations occurred with equal probability of 12.5 each. Clear MMNm was obtained for most changes in both groups, despite the 50 deviance level, with a larger amplitude in musicians than in controls. The response pattern was consistent across groups, with larger MMNm for deviants in the high voice than in the low voice, and larger MMNm for in-key than out-of-key changes, despite better behavioral performance for out-of-key changes. The results suggest that melodic information in each voice in polyphonic music is encoded in the sensory memory trace, that the higher voice is more salient than the lower, and that tonality may be processed primarily at cognitive stages subsequent to MMN generation.

Aging in Binaural Hearing Begins in Mid-Life: Evidence from Cortical Auditory-Evoked Responses to Changes in Interaural Phase

Older adults often have difficulty understanding speech in a noisy environment or with multiple speakers. In such situations, binaural hearing improves the signal-to-noise ratio. How does this binaural advantage change with increasing age? Using magnetoencephalography, we recorded cortical activity evoked by changes in interaural phase differences of amplitude-modulated tones. These responses occurred for frequencies up to 1225 Hz in young subjects but only up to 940 Hz in middle-aged and 760 Hz in older adults. Behavioral thresholds also decreased with increasing age but were more variable, likely because some older adults make effective use of compensatory mechanisms. The reduced frequency range for binaural hearing became significant in middle age, before decline in hearing sensation and the morphology of cortical responses, which became apparent only in the older subjects. This study provides evidence from human physiological data for the early onset of biological aging in binaural hearing.

Endogenous Neuromagnetic Activity for Mental Hierarchy of Timing

The frontal-striatal circuits, the cerebellum, and motor cortices play crucial roles in processing timing information on second to millisecond scales. However, little is known about the physiological mechanism underlying humans preference to robustly encode a sequence of time intervals into a mental hierarchy of temporal units called meter. This is especially salient in music: temporal patterns are typically interpreted as integer multiples of a basic unit (i.e., the beat) and accommodated into a global context such as march or waltz. With magnetoencephalography and spatial-filtering source analysis, we demonstrated that the time courses of neural activities index a subjectively induced meter context. Auditory evoked responses from hippocampus, basal ganglia, and auditory and association cortices showed a significant contrast between march and waltz metric conditions during listening to identical click stimuli. Specifically, the right hippocampus was activated differentially at 80 ms to the march downbeat (the count one) and ∼250 ms to the waltz downbeat. In contrast, basal ganglia showed a larger 80 ms peak for march downbeat than waltz. The metric contrast was also expressed in long-latency responses in the right temporal lobe. These findings suggest that anticipatory processes in the hippocampal memory system and temporal computation mechanism in the basal ganglia circuits facilitate endogenous activities in auditory and association cortices through feedback loops. The close interaction of auditory, motor, and limbic systems suggests a distributed network for metric organization in temporal processing and its relevance for musical behavior.

Beta-Band Oscillations Represent Auditory Beat and Its Metrical Hierarchy in Perception and Imagery

Dancing to music involves synchronized movements, which can be at the basic beat level or higher hierarchical metrical levels, as in a march (groups of two basic beats, one–two–one–two …) or waltz (groups of three basic beats, one–two–three–one–two–three …). Our previous human magnetoencephalography studies revealed that the subjective sense of meter influences auditory evoked responses phase locked to the stimulus. Moreover, the timing of metronome clicks was represented in periodic modulation of induced (non-phase locked) β-band (13–30 Hz) oscillation in bilateral auditory and sensorimotor cortices. Here, we further examine whether acoustically accented and subjectively imagined metric processing in march and waltz contexts during listening to isochronous beats were reflected in neuromagnetic β-band activity recorded from young adult musicians. First, we replicated previous findings of beat-related β-power decrease at 200 ms after the beat followed by a predictive increase toward the onset of the next beat. Second, we showed that the β decrease was significantly influenced by the metrical structure, as reflected by differences across beat type for both perception and imagery conditions. Specifically, the β-power decrease associated with imagined downbeats (the count “one”) was larger than that for both the upbeat (preceding the count “one”) in the march, and for the middle beat in the waltz. Moreover, beamformer source analysis for the whole brain volume revealed that the metric contrasts involved auditory and sensorimotor cortices; frontal, parietal, and inferior temporal lobes; and cerebellum. We suggest that the observed β-band activities reflect a translation of timing information to auditory–motor coordination. SIGNIFICANCE STATEMENT With magnetoencephalography, we examined β-band oscillatory activities around 20 Hz while participants listened to metronome beats and imagined musical meters such as a march and waltz. We demonstrated that β-band event-related desynchronization in the auditory cortex differentiates between beat positions, specifically between downbeats and the following beat. This is the first demonstration of β-band oscillations related to hierarchical and internalized timing information. Moreover, the meter representation in the β oscillations was widespread across the brain, including sensorimotor and premotor cortices, parietal lobe, and cerebellum. The results extend current understanding of the role of β oscillations in neural processing of predictive timing.

Neural Representation of Transposed Melody in Infants at 6 Months of Age

We examined adults’ and 6‐month‐old infants’ event‐related potentials in response to occasional changes (deviants) in a 4‐note melody presented at different pitch levels from trial to trial. In both groups, responses to standard and deviant stimuli differed significantly; however, adults produced a typical mismatch negativity (MMN), whereas 6‐month‐old infants exhibited a slow positive wave. We conclude that 6‐month‐old infants, like adults, encode melodic information in terms of relative pitch distances, but that the underlying cortical activity differs significantly from that of adults.

Beat-induced fluctuations in auditory cortical beta-band activity: using EEG to measure age-related changes.

People readily extract regularity in rhythmic auditory patterns, enabling prediction of the onset of the next beat. Recent magnetoencephalography (MEG) research suggests that such prediction is reflected by the entrainment of oscillatory networks in the brain to the tempo of the sequence. In particular, induced beta-band oscillatory activity from auditory cortex decreases after each beat onset and rebounds prior to the onset of the next beat across tempi in a predictive manner. The objective of the present study was to examine the development of such oscillatory activity by comparing electroencephalography (EEG) measures of beta-band fluctuations in 7-year-old children to adults. EEG was recorded while participants listened passively to isochronous tone sequences at three tempi (390, 585, and 780 ms for onset-to-onset interval). In adults, induced power in the high beta-band (20–25 Hz) decreased after each tone onset and rebounded prior to the onset of the next tone across tempo conditions, consistent with MEG findings. In children, a similar pattern was measured in the two slower tempo conditions, but was weaker in the fastest condition. The results indicate that the beta-band timing network works similarly in children, although there are age-related changes in consistency and the tempo range over which it operates.

Synchronization of beta and gamma oscillations in the somatosensory evoked neuromagnetic steady-state response

The sensory evoked neuromagnetic response consists of superimposition of an immediately stimulus-driven component and induced changes in the autonomous brain activity, each having distinct functional relevance. Commonly, the strength of phase locking in neural activities has been used to differentiate the different responses. The steady-state response is a strong oscillatory neural activity, which is evoked with rhythmic stimulation, and provides an effective tool to investigate oscillatory brain networks. In this case, both the sensory response and intrinsic activity, representing higher order processes, are highly synchronized to the stimulus. In this study we hypothesized that temporal dynamics of oscillatory activities would characterize the differences between the two types of activities and that beta and gamma oscillations are differently involved in this distinction. We used magnetoencephalography (MEG) for studying how ongoing steady-state responses elicited by a 20-Hz vibro-tactile stimulus to the right index finger were affected by a concurrent isolated touch stimulus to the same hand ring finger. SI source activity showed oscillations at multiples of 20 Hz with characteristic differences in the beta band and the gamma band. The response amplitudes were largest at 20 Hz (beta) and significantly reduced at 40 Hz and 60 Hz (gamma), although synchronization strength, indicated by inter-trial coherence (ITC), did not substantially differ between 20 Hz and 40 Hz. Moreover, the beta oscillations showed a fast onset, whereas the amplitude of gamma oscillations increased slowly and reached the steady state 400 ms after onset of the vibration stimulus. Most importantly, the pulse stimuli interacted only with gamma oscillations in a way that gamma oscillations decreased immediately after the concurrent stimulus onset and recovered slowly, resembling the initial slope. Such time course of gamma oscillations is similar to our previous observations in the auditory system. The time constant is in line with the time required for conscious perception of the sensory stimulus. Based on the observed different spectro-temporal dynamics, we propose that while beta activities likely relate to independent representation of the sensory input, gamma oscillation likely relates to binding of sensory information for higher order processing.