Macaque monkeys and humans sample temporal regularities in the acoustic environment.

Many animal species show comparable abilities to detect basic rhythms and produce rhythmic behavior. Yet, the capacities to process complex rhythms and synchronize rhythmic behavior appear to be species-specific: vocal learning animals can, but some primates might not. This discrepancy is of high interest as there is a putative link between rhythm processing and the development of sophisticated sensorimotor behavior in humans. Do our closest ancestors show comparable endogenous dispositions to sample the acoustic environment in the absence of task instructions and training? We recorded EEG from macaque monkeys and humans while they passively listened to isochronous equitone sequences. Individual- and trial-level analyses showed that macaque monkeys' and humans' delta-band neural oscillations encoded and tracked the timing of auditory events. Further, mu- (8-15 Hz) and beta-band (12-20 Hz) oscillations revealed the superimposition of varied accentuation patterns on a subset of trials. These observations suggest convergence in the encoding and dynamic attending of temporal regularities in the acoustic environment, bridging a gap in the phylogenesis of rhythm cognition.

Sensitivity to syllable stress regularities in externally but not self-triggered speech in Dutch.

Several theories of predictive processing propose reduced sensory and neural responses to anticipated events. Support comes from magnetoencephalography/electroencephalography (M/EEG) studies, showing reduced auditory N1 and P2 responses to self-generated compared to externally generated events, or when the timing and form of stimuli are more predictable. The current study examined the sensitivity of N1 and P2 responses to statistical speech regularities. We employed a motor-to-auditory paradigm comparing event-related potential (ERP) responses to externally and self-triggered pseudowords. Participants were presented with a cue indicating which button to press (motor-auditory condition) or which pseudoword would be presented (auditory-only condition). Stimuli consisted of the participant's own voice uttering pseudowords that varied in phonotactic probability and syllable stress. We expected to see N1 and P2 suppression for self-triggered stimuli, with greater suppression effects for more predictable features such as high phonotactic probability and first-syllable stress in pseudowords. In a temporal principal component analysis (PCA), we observed an interaction between syllable stress and condition for the N1, where second-syllable stress items elicited a larger N1 than first-syllable stress items, but only for externally generated stimuli. We further observed an effect of syllable stress on the P2, where first-syllable stress items elicited a larger P2. Strikingly, we did not observe motor-induced suppression for self-triggered stimuli for either the N1 or P2 component, likely due to the temporal predictability of the stimulus onset in both conditions. Taking into account previous findings, the current results suggest that sensitivity to syllable stress regularities depends on task demands.

Sensory gating functions of the auditory thalamus: Adaptation and modulations through noise-exposure and high-frequency stimulation in rats.

The medial geniculate body (MGB) of the thalamus is an obligatory relay for auditory processing. A breakdown of adaptive filtering and sensory gating at this level may lead to multiple auditory dysfunctions, while high-frequency stimulation (HFS) of the MGB might mitigate aberrant sensory gating. To further investigate the sensory gating functions of the MGB, this study (i) recorded electrophysiological evoked potentials in response to continuous auditory stimulation, and (ii) assessed the effect of MGB HFS on these responses in noise-exposed and control animals. Pure-tone sequences were presented to assess differential sensory gating functions associated with stimulus pitch, grouping (pairing), and temporal regularity. Evoked potentials were recorded from the MGB and acquired before and after HFS (100 Hz). All animals (unexposed and noise-exposed, pre- and post-HFS) showed gating for pitch and grouping. Unexposed animals also showed gating for temporal regularity not found in noise-exposed animals. Moreover, only noise-exposed animals showed restoration comparable to the typical EP amplitude suppression following MGB HFS. The current findings confirm adaptive thalamic sensory gating based on different sound characteristics and provide evidence that temporal regularity affects MGB auditory signaling.

Sleeping with time in mind? A literature review and a proposal for a screening questionnaire on self-awakening.

Some people report being able to spontaneously "time" the end of their sleep. This ability to self-awaken challenges the idea of sleep as a passive cognitive state. Yet, current evidence on this phenomenon is limited, partly because of the varied definitions of self-awakening and experimental approaches used to study it. Here, we provide a review of the literature on self-awakening. Our aim is to i) contextualise the phenomenon, ii) propose an operating definition, and iii) summarise the scientific approaches used so far. The literature review identified 17 studies on self-awakening. Most of them adopted an objective sleep evaluation (76%), targeted nocturnal sleep (76%), and used a single criterion to define the success of awakening (82%); for most studies, this corresponded to awakening occurring in a time window of 30 minutes around the expected awakening time. Out of 715 total participants, 125 (17%) reported to be self-awakeners, with an average age of 23.24 years and a slight predominance of males compared to females. These results reveal self-awakening as a relatively rare phenomenon. To facilitate the study of self-awakening, and based on the results of the literature review, we propose a quick paper-and-pencil screening questionnaire for self-awakeners and provide an initial validation for it. Taken together, the combined results of the literature review and the proposed questionnaire help in characterising a theoretical framework for self-awakenings, while providing a useful tool and empirical suggestions for future experimental studies, which should ideally employ objective measurements.

Magnetoencephalography recordings reveal the spatiotemporal dynamics of recognition memory for complex versus simple auditory sequences.

Auditory recognition is a crucial cognitive process that relies on the organization of single elements over time. However, little is known about the spatiotemporal dynamics underlying the conscious recognition of auditory sequences varying in complexity. To study this, we asked 71 participants to learn and recognize simple tonal musical sequences and matched complex atonal sequences while their brain activity was recorded using magnetoencephalography (MEG). Results reveal qualitative changes in neural activity dependent on stimulus complexity: recognition of tonal sequences engages hippocampal and cingulate areas, whereas recognition of atonal sequences mainly activates the auditory processing network. Our findings reveal the involvement of a cortico-subcortical brain network for auditory recognition and support the idea that stimulus complexity qualitatively alters the neural pathways of recognition memory.

Left Motor δ Oscillations Reflect Asynchrony Detection in Multisensory Speech Perception.

During multisensory speech perception, slow δ oscillations (∼1-3 Hz) in the listener's brain synchronize with the speech signal, likely engaging in speech signal decomposition. Notable fluctuations in the speech amplitude envelope, resounding speaker prosody, temporally align with articulatory and body gestures and both provide complementary sensations that temporally structure speech. Further, δ oscillations in the left motor cortex seem to align with speech and musical beats, suggesting their possible role in the temporal structuring of (quasi)-rhythmic stimulation. We extended the role of δ oscillations to audiovisual asynchrony detection as a test case of the temporal analysis of multisensory prosody fluctuations in speech. We recorded Electroencephalograph (EEG) responses in an audiovisual asynchrony detection task while participants watched videos of a speaker. We filtered the speech signal to remove verbal content and examined how visual and auditory prosodic features temporally (mis-)align. Results confirm (1) that participants accurately detected audiovisual asynchrony, and (2) increased δ power in the left motor cortex in response to audiovisual asynchrony. The difference of δ power between asynchronous and synchronous conditions predicted behavioral performance, and (3) decreased δ-ß coupling in the left motor cortex when listeners could not accurately map visual and auditory prosodies. Finally, both behavioral and neurophysiological evidence was altered when a speaker's face was degraded by a visual mask. Together, these findings suggest that motor δ oscillations support asynchrony detection of multisensory prosodic fluctuation in speech.SIGNIFICANCE STATEMENT Speech perception is facilitated by regular prosodic fluctuations that temporally structure the auditory signal. Auditory speech processing involves the left motor cortex and associated δ oscillations. However, visual prosody (i.e., a speaker's body movements) complements auditory prosody, and it is unclear how the brain temporally analyses different prosodic features in multisensory speech perception. We combined an audiovisual asynchrony detection task with electroencephalographic (EEG) recordings to investigate how δ oscillations support the temporal analysis of multisensory speech. Results confirmed that asynchrony detection of visual and auditory prosodies leads to increased δ power in left motor cortex and correlates with performance. We conclude that δ oscillations are invoked in an effort to resolve denoted temporal asynchrony in multisensory speech perception.

About time: Ageing influences neural markers of temporal predictability.

Timing abilities help organizing the temporal structure of events but are known to change systematically with age. Yet, how the neuronal signature of temporal predictability changes across the age span remains unclear. Younger (n = 21; 23.1 years) and older adults (n = 21; 68.5 years) performed an auditory oddball task, consisting of isochronous and random sound sequences. Results confirm an altered P50 response in the older compared to younger participants. P50 amplitudes differed between the isochronous and random temporal structures in younger, and for P200 in the older group. These results suggest less efficient sensory gating in older adults in both isochronous and random auditory sequences. N100 amplitudes were more negative for deviant tones. P300 amplitudes were parietally enhanced in younger, but not in older adults. In younger participants, the P50 results confirm that this component marks temporal predictability, indicating sensitive gating of temporally regular sound sequences.

Auditory thalamus dysfunction and pathophysiology in tinnitus: a predictive network hypothesis.

Tinnitus is the perception of a 'ringing' sound without an acoustic source. It is generally accepted that tinnitus develops after peripheral hearing loss and is associated with altered auditory processing. The thalamus is a crucial relay in the underlying pathways that actively shapes processing of auditory signals before the respective information reaches the cerebral cortex. Here, we review animal and human evidence to define thalamic function in tinnitus. Overall increased spontaneous firing patterns and altered coherence between the thalamic medial geniculate body (MGB) and auditory cortices is observed in animal models of tinnitus. It is likely that the functional connectivity between the MGB and primary and secondary auditory cortices is reduced in humans. Conversely, there are indications for increased connectivity between the MGB and several areas in the cingulate cortex and posterior cerebellar regions, as well as variability in connectivity between the MGB and frontal areas regarding laterality and orientation in the inferior, medial and superior frontal gyrus. We suggest that these changes affect adaptive sensory gating of temporal and spectral sound features along the auditory pathway, reflecting dysfunction in an extensive thalamo-cortical network implicated in predictive temporal adaptation to the auditory environment. Modulation of temporal characteristics of input signals might hence factor into a thalamo-cortical dysrhythmia profile of tinnitus, but could ultimately also establish new directions for treatment options for persons with tinnitus.

Dynamic acoustic salience evokes motor responses.

Audio-motor integration is currently viewed as a predictive process in which the brain simulates upcoming sounds based on voluntary actions. This perspective does not consider how our auditory environment may trigger involuntary action in the absence of prediction. We address this issue by examining the relationship between acoustic salience and involuntary motor responses. We investigate how acoustic features in music contribute to the perception of salience, and whether those features trigger involuntary peripheral motor responses. Participants with little-to-no musical training listened to musical excerpts once while remaining still during the recording of their muscle activity with surface electromyography (sEMG), and again while they continuously rated perceived salience within the music using a slider. We show cross-correlations between 1) salience ratings and acoustic features, 2) acoustic features and spontaneous muscle activity, and 3) salience ratings and spontaneous muscle activity. Amplitude, intensity, and spectral centroid were perceived as the most salient features in music, and fluctuations in these features evoked involuntary peripheral muscle responses. Our results suggest an involuntary mechanism for audio-motor integration, which may rely on brainstem-spinal or brainstem-cerebellar-spinal pathways. Based on these results, we argue that a new framework is needed to explain the full range of human sensorimotor capabilities. This goal can be achieved by considering how predictive and reactive audio-motor integration mechanisms could operate independently or interactively to optimize human behavior.

ERP mismatch response to phonological and temporal regularities in speech.

Predictions of our sensory environment facilitate perception across domains. During speech perception, formal and temporal predictions may be made for phonotactic probability and syllable stress patterns, respectively, contributing to the efficient processing of speech input. The current experiment employed a passive EEG oddball paradigm to probe the neurophysiological processes underlying temporal and formal predictions simultaneously. The component of interest, the mismatch negativity (MMN), is considered a marker for experience-dependent change detection, where its timing and amplitude are indicative of the perceptual system's sensitivity to presented stimuli. We hypothesized that more predictable stimuli (i.e. high phonotactic probability and first syllable stress) would facilitate change detection, indexed by shorter peak latencies or greater peak amplitudes of the MMN. This hypothesis was confirmed for phonotactic probability: high phonotactic probability deviants elicited an earlier MMN than low phonotactic probability deviants. We do not observe a significant modulation of the MMN to variations in syllable stress. Our findings confirm that speech perception is shaped by formal and temporal predictability. This paradigm may be useful to investigate the contribution of implicit processing of statistical regularities during (a)typical language development.

When temporal prediction errs: ERP responses to delayed action-feedback onset.

Sensory suppression effects observed in electroencephalography (EEG) index successful predictions of the type and timing of self-generated sensory feedback. However, it is unclear how precise the timing prediction of sensory feedback is, and how temporal delays between an action and its sensory feedback affect perception. The current study investigated how prediction errors induced by delaying tone onset times affect the processing of sensory feedback in audition. Participants listened to self-generated (via button press) or externally generated tones. Self-generated tones were presented either without or with various delays (50, 100, or 250 ms; in 30% of trials). Comparing listening to externally generated and self-generated tones resulted in action-related P50 amplitude suppression to tones presented immediately or 100 ms after the button press. Subsequent ERP responses became more sensitive to the type of delay. Whereas the comparison of actual and predicted sensory feedback (N1) tolerated temporal uncertainty up to 100 ms, P2 suppression was modulated by delay in a graded manner: suppression decreased with an increase in sensory feedback delay. Self-generated tones occurring 250 ms after the button press additionally elicited an enhanced N2 response. These findings suggest functionally dissociable processes within the forward model that are affected by the timing of sensory feedback to self-action: relative tolerance of temporal delay in the P50 and N1, confirming previous results, but increased sensitivity in the P2. Further, they indicate that temporal prediction errors are treated differently by the auditory system: only delays that occurred after a temporal integration window (∼100 ms) impact the conscious detection of altered sensory feedback.

Beta power encodes contextual estimates of temporal event probability in the human brain.

To prepare for an impending event of unknown temporal distribution, humans internally increase the perceived probability of event onset as time elapses. This effect is termed the hazard rate of events. We tested how the neural encoding of hazard rate changes by providing human participants with prior information on temporal event probability. We recorded behavioral and electroencephalographic (EEG) data while participants listened to continuously repeating five-tone sequences, composed of four standard tones followed by a non-target deviant tone, delivered at slow (1.6 Hz) or fast (4 Hz) rates. The task was to detect a rare target tone, which equiprobably appeared at either position two, three or four of the repeating sequence. In this design, potential target position acts as a proxy for elapsed time. For participants uninformed about the target's distribution, elapsed time to uncertain target onset increased response speed, displaying a significant hazard rate effect at both slow and fast stimulus rates. However, only in fast sequences did prior information about the target's temporal distribution interact with elapsed time, suppressing the hazard rate. Importantly, in the fast, uninformed condition pre-stimulus power synchronization in the beta band (Beta 1, 15-19 Hz) predicted the hazard rate of response times. Prior information suppressed pre-stimulus power synchronization in the same band, while still significantly predicting response times. We conclude that Beta 1 power does not simply encode the hazard rate, but-more generally-internal estimates of temporal event probability based upon contextual information.

Laughter catches attention!

In social interactions, emotionally salient and sudden changes in vocal expressions attract attention. However, only a few studies examined how emotion and attention interact in voice processing. We investigated neutral, happy (laughs) and angry (growls) vocalizations in a modified oddball task. Participants silently counted the targets in each block and rated the valence and arousal of the vocalizations. A combined event-related potential and time-frequency analysis focused on the P3 and pre-stimulus alpha power to capture attention effects in response to unexpected events. Whereas an early differentiation between emotionally salient and neutral vocalizations was reflected in the P3a response, the P3b was selectively enhanced for happy voices. The P3b modulation was predicted by pre-stimulus frontal alpha desynchronization, and by the perceived pleasantness of the targets. These findings indicate that vocal emotions may be differently processed based on task relevance and valence. Increased anticipation and attention to positive vocal cues (laughter) may reflect their high social relevance.

Is laughter a better vocal change detector than a growl?

The capacity to predict what should happen next and to minimize any discrepancy between an expected and an actual sensory input (prediction error) is a central aspect of perception. Particularly in vocal communication, the effective prediction of an auditory input that informs the listener about the emotionality of a speaker is critical. What is currently unknown is how the perceived valence of an emotional vocalization affects the capacity to predict and detect a change in the auditory input. This question was probed in a combined event-related potential (ERP) and time-frequency analysis approach. Specifically, we examined the brain response to standards (Repetition Positivity) and to deviants (Mismatch Negativity - MMN), as well as the anticipatory response to the vocal sounds (pre-stimulus beta oscillatory power). Short neutral, happy (laughter), and angry (growls) vocalizations were presented both as standard and deviant stimuli in a passive oddball listening task while participants watched a silent movie and were instructed to ignore the vocalizations. MMN amplitude was increased for happy compared to neutral and angry vocalizations. The Repetition Positivity was enhanced for happy standard vocalizations. Induced pre-stimulus upper beta power was increased for happy vocalizations, and predicted the modulation of the standard Repetition Positivity. These findings indicate enhanced sensory prediction for positive vocalizations such as laughter. Together, the results suggest that positive vocalizations are more effective predictors in social communication than angry and neutral ones, possibly due to their high social significance.

Gait improvement via rhythmic stimulation in Parkinson's disease is linked to rhythmic skills.

Training based on rhythmic auditory stimulation (RAS) can improve gait in patients with idiopathic Parkinson's disease (IPD). Patients typically walk faster and exhibit greater stride length after RAS. However, this effect is highly variable among patients, with some exhibiting little or no response to the intervention. These individual differences may depend on patients' ability to synchronize their movements to a beat. To test this possibility, 14 IPD patients were submitted to RAS for four weeks, in which they walked to music with an embedded metronome. Before and after the training, patients' synchronization was assessed with auditory paced hand tapping and walking to auditory cues. Patients increased gait speed and stride length in non-cued gait after training. However, individual differences were apparent as some patients showed a positive response to RAS and others, either no response, or a negative response. A positive response to RAS was predicted by the synchronization performance in hand tapping and gait tasks. More severe gait impairment, low synchronization variability, and a prompt response to a stimulation change foster a positive response to RAS training. Thus, sensorimotor timing skills underpinning the synchronization of steps to an auditory cue may allow predicting the success of RAS in IPD.

Uncertainty and expectancy deviations require cortico-subcortical cooperation.

In a dynamic and uncertain environment it is beneficial to learn the causal structure of the environment in order to minimize uncertainty. This requires determining estimates of probable outcomes, which will guide expectations about incoming information. One key factor in this learning process is to detect whether an unexpected event constitutes a low probability, but valid outcome, or an outright error. The present 7T-fMRI study investigated the role of subcortical structures in regulating this probabilistic inferential learning process. A new task was designed, in which participants learned to calculate the value, and therefore to anticipate the outcome of different visual sequences. Three types of sequences provided unambiguous, ambiguous, and incongruent contextual evidence and each sequence had two outcomes, which differed in their probability of occurrence. We hypothesized that subcortical regions are necessary when expectations are violated, and that their involvement will depend on the nature of the unexpected event. The results show increased dorsomedial striatal and thalamic activation for less probable sequences; in addition, ambiguous sequences also display larger activation in the red nuclei. Incongruent sequences displayed a pattern of subcortical activation restricted to the dorsolateral and the posterior dorsomedial striatum. These results confirm that different subcortical structures regulate uncertainty and expectancy deviations; this is crucial not only for learning to predict events in the environment, but also for flexible cognitive control in general.

Striatal contributions to sensory timing: Voxel-based lesion mapping of electrophysiological markers.

To achieve precise timing, the brain needs to establish a representation of the temporal structure of sensory input and use this information to generate timely responses. These operations engage the basal ganglia. Current research in this direction is limited by reliance on animal models, motor and/or offline tasks, small sample sizes, the low temporal resolution of functional magnetic resonance imaging, and the study of progressive neurodegeneration. Here, we combine the excellent temporal resolution of electrophysiological potentials with the high spatial resolution of structural neuroimaging to investigate basal ganglia contributions to sensory timing. Chronic-stage lesion patients and healthy controls listened to pure-tone sequences differing exclusively in temporal regularity. Event-related potentials (ERPs) indicate a selective indifference against this manipulation in patients, attributable to the striatal part of the basal ganglia on the basis of a lesion-mapping approach. These findings provide evidence for a crucial contribution of the basal ganglia to basic sensory functioning.

Basal ganglia contribution to rule expectancy and temporal predictability in speech.

The current work set out to answer three questions: (1) Are reported syntactic deficits in patients with structural damage to the basal ganglia (BG) in the cortico-striato-thalamo-cortical systems (CSTCS) the result of a syntax specific computational deficit or are they potentially a consequence of a generalized timing deficit? (2) Do BG patients suffer from a simple beat perception deficit in speech comparable to the one reported in music? (3) Can regular speech meter (i.e., a pattern of beats induced by the regular alteration of stressed and unstressed syllable accents) ameliorate the computation of syntactically marked information by making speech events temporally predictable and salient? The latter 'remediation' hypothesis would predict that when speech events (i.e., those that are syntactically marked) are metrically aligned to the syllabic accent structure, the computation of syntactic information is facilitated or in the case of patients ameliorated. During continuous EEG measurement nineteen patients with focal BG lesions and matched healthy controls listened to metrically regular and syntactically well-formed sentences and metrically well-formed sentences that either violated syntactic expectancy, metrical expectancy, or both. While healthy controls showed an expected P600 response in the event-related brain potential (ERP) to all expectancy violations, BG patients showed overall comparable P600 responses to all, but the metrical expectancy violation. These results confirm that (1) BG patients suffer from a simple beat perception deficit in speech and (2) regular speech meter ameliorates the computation of syntactically marked information in the speech signal. We propose that a domain general sensorimotor cerebello-thalamo-cortical system (CTCS), involved in event-based temporal processing, engages in the remediation of dysfunctional cortico-striato-thalamo-cortical timing that affects the timely computation of linguistic (i.e., syntax) information in the speech signal.

Effects of musically cued gait training in Parkinson's disease: beyond a motor benefit.

Auditory stimulation via rhythmic cues can be used successfully in the rehabilitation of motor function in patients with motor disorders. A prototypical example is provided by dysfunctional gait in patients with idiopathic Parkinson's disease (PD). Coupling steps to external rhythmic cues (the beat of music or the sounds of a metronome) leads to long-term motor improvements, such as increased walking speed and greater stride length. These effects are likely to be underpinned by compensatory brain mechanisms involving cerebellar-thalamocortical networks. Because these areas are also involved in perceptual and motor timing, parallel improvement in timing tasks is expected in PD beyond purely motor benefits. In keeping with this idea, we report here recent behavioral data showing beneficial effects of musically cued gait training (MCGT) on gait performance (i.e., increased stride length and speed), perceptual timing (e.g., discriminating stimulus durations), and sensorimotor timing abilities (i.e., in paced tapping tasks) in PD patients. Particular attention is paid to individual differences in timing abilities in PD, thus paving the ground for an individualized MCGT-based therapy.

Temporal dynamics of contingency extraction from tonal and verbal auditory sequences.

Consecutive sound events are often to some degree predictive of each other. Here we investigated the brain's capacity to detect contingencies between consecutive sounds by means of electroencephalography (EEG) during passive listening. Contingencies were embedded either within tonal or verbal stimuli. Contingency extraction was measured indirectly via the elicitation of the mismatch negativity (MMN) component of the event-related potential (ERP) by contingency violations. MMN results indicate that structurally identical forms of predictability can be extracted from both tonal and verbal stimuli. We also found similar generators to underlie the processing of contingency violations across stimulus types, as well as similar performance in an active-listening follow-up test. However, the process of passive contingency extraction was considerably slower (twice as many rule exemplars were needed) for verbal than for tonal stimuli These results suggest caution in transferring findings on complex predictive regularity processing obtained with tonal stimuli directly to the speech domain.