Search

Self-generated auditory input is perceived less loudly than the same sounds generated externally. The existence of this phenomenon, called Sensory Attenuation (SA), has been studied for decades and is often explained by motor-based forward models. Recent developments in the research of SA, however, challenge these models. We review the current state of knowledge regarding theoretical implications about the significance of Sensory Attenuation and its role in human behavior and functioning. Focusing on behavioral and electrophysiological results in the auditory domain, we provide an overview of the characteristics and limitations of existing SA paradigms and highlight the problem of isolating SA from other predictive mechanisms. Finally, we explore different hypotheses attempting to explain heterogeneous empirical findings, and the impact of the Predictive Coding Framework in this research area.

Rhythmic stimulation is a powerful tool to improve temporal prediction and parsing of the auditory signal. However, for long duration of stimulation, the rhythmic and repetitive aspects of music have often been associated to a trance state. In this study we conceived an auditory monitoring task that allows tracking changes of psychophysical auditory thresholds. Participants performed the task while listening to rhythmically regular and an irregular (scrambled but spectrally identical) music that were presented with an intermittent (short) and continuous (long) type of stimulation. Results show that psychophysical auditory thresholds increase following a Continuous versus Intermittent stimulation and this is accompanied by a reduction of the amplitude of two event-related potentials to target stimuli. These effects are larger with regular music, thus do not simply derive from the duration of stimulation. Interestingly, they seem to be related to a frequency selective neural coupling as well as an increase of network connectivity in the alpha band between frontal and central regions. Our study shows that the idea that rhythmic presentation of sensory stimuli facilitates perception might be limited to short streams, while long, highly regular, repetitive and strongly engaging streams may have an opposite perceptual impact.

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.

Behavioral and brain rhythms in the millisecond-to-second range are central in human music, speech, and movement. A comparative approach can further our understanding of the evolution of rhythm processing by identifying behavioral and neural similarities and differences across cognitive domains and across animal species. We provide an overview of research into rhythm cognition in music, speech, and animal communication. Rhythm has received considerable attention within each individual field, but to date, little integration. This review article on rhythm processing incorporates and extends existing ideas on temporal processing in speech and music and offers suggestions about the neural, biological, and evolutionary bases of human abilities in these domains.

International audience; While the benefit of temporal predictability on sensorimotor processing is well established, it is still unknown whether this is due to efficient execution of an appropriate response and/or inhibition of an inappropriate one. To answer this question, we examined the effects of temporal predictability in tasks that required selective (Simon task) or global (Stop-signal task) inhibitory control of prepotent responses. We manipulated temporal expectation by presenting cues that either predicted (temporal cues) or not (neutral cues) when the target would appear. In the Simon task, performance was better when target location (left/right) was compatible with the hand of response and performance was improved further still if targets were temporally cued. However, Conditional Accuracy Functions revealed that temporal predictability selectively increased the number of fast, impulsive errors. Temporal cueing had no effect on selective response inhibition, as measured by the dynamics of the interference effect (delta plots) in the Simon task. By contrast, in the Stop-signal task, Stop-signal reaction time, a covert measure of a more global form of response inhibition, was significantly longer in temporally predictive trials. Therefore, when the time of target onset could be predicted in advance, it was harder to stop the impulse to respond to the target. Collectively, our results indicate that temporal cueing compounded the interfering effects of a prepotent response on task performance. We suggest that although temporal predictability enhances activation of task-relevant responses, it impairs inhibition of prepotent responses.

Music can be thought of as a dynamic path over time. In most cases, the rhythmic structure of this path, such as specific sequences of strong and weak beats or recurring patterns, allows us to predict what and particularly when sounds are going to happen. Without this ability we would not be able to entrain body movements to music, like we do when we dance. By combining EEG and behavioral measures, the current study provides evidence illustrating the importance of ongoing neural oscillations at beat-related frequencies – i.e., neural entrainment – for tracking and predicting musical rhythms. Participants (13 musicians and 13 non-musicians) listened to drum rhythms that switched from a quadruple rhythm to a 3-over-4 polyrhythm. After a silent period of approximately two to three seconds, participants had to decide whether a target stimulus was presented on time with the triple beat of the polyrhythm, too early, or too late. Results showed that neural oscillations reflected the rhythmic structure of both the simple quadruple rhythm and the more complex polyrhythm with no differences between musicians and non-musicians. During silent periods, the observation of time-frequency plots and more commonly used frequency spectra analyses suggest that beat-related neural oscillations were more pronounced in musicians compared to non-musicians. Neural oscillations during silent periods are not driven by an external input and therefore are thought to reflect top-down controlled endogenous neural entrainment. The functional relevance of endogenous neural entrainment was demonstrated by a positive correlation between the amplitude of task-relevant neural oscillations during silent periods and the number of correctly identified target stimuli. In sum, our findings add to the evidence supporting the neural resonance theory of pulse and meter. Furthermore, they indicate that beat-related top-down controlled neural oscillations can exist without external stimulation and suggest that those endogenous oscillations are strengthened by musical expertise. Finally, this study shows that the analysis of neural oscillations can be a useful tool to assess how we perceive and process complex auditory stimuli such as polyrhythms.

Temporal structure in the environment often has predictive value for anticipating the occurrence of forthcoming events. In this study we investigated the influence of two types of predictive temporal information on the perception of near-threshold auditory stimuli: 1) intrinsic temporal rhythmicity within an auditory stimulus stream and 2) temporally-predictive visual cues. We hypothesized that combining predictive temporal information within- and across-modality should decrease the threshold at which sounds are detected, beyond the advantage provided by each information source alone. Two experiments were conducted in which participants had to detect tones in noise. Tones were presented in either rhythmic or random sequences and were preceded by a temporally predictive visual signal in half of the trials. We show that detection intensities are lower for rhythmic (vs. random) and audiovisual (vs. auditory-only) presentation, independent from response bias, and that this effect is even greater for rhythmic audiovisual presentation. These results suggest that both types of temporal information are used to optimally process sounds that occur at expected points in time (resulting in enhanced detection), and that multiple temporal cues are combined to improve temporal estimates. Our findings underscore the flexibility and proactivity of the perceptual system which uses within- and across-modality temporal cues to anticipate upcoming events and process them optimally. (C) 2014 Elsevier Ltd. All rights reserved.

Moving to a speech rhythm can enhance verbal processing in the listener by increasing temporal expectancies (Falk and Dalla Bella, 2016). Here we tested whether this hypothesis holds for prosodically diverse languages such as German (a lexical stress-language) and French (a non-stress language). Moreover, we examined the relation between motor performance and the benefits for verbal processing as a function of language. Sixty-four participants, 32 German and 32 French native speakers detected subtle word changes in accented positions in metrically structured sentences to which they previously tapped with their index finger. Before each sentence, they were cued by a metronome to tap either congruently (i.e., to accented syllables) or incongruently (i.e., to non-accented parts) to the following speech stimulus. Both French and German speakers detected words better when cued to tap congruently compared to incongruent tapping. Detection performance was predicted by participants' motor performance in the non-verbal cueing phase. Moreover, tapping rate while participants tapped to speech predicted detection differently for the two language groups, in particular in the incongruent tapping condition. We discuss our findings in light of the rhythmic differences of both languages and with respect to recent theories of expectancy-driven and multisensory speech processing.

International audience; Recent advances in neuroscience have brought a great focus on how the auditory cortex tracks speech at certain time scales corresponding to pre-lexical speech units in order to achieve comprehension. In particular, it has been claimed that it is the syllabic rhythm to which slow neural oscillations in the auditory cortex entrain in order to chunk the speech stream into smaller informational units. However, the terms “syllable” and “rhythm” have been treated quite loosely in the current literature. We revisit classic approaches to show that both concepts do not necessarily have an acoustic or phonetic counterpart, which could be directly extracted by neural processes. We would like to suggest that the syllabic rhythm could emerge at the intersection of acoustic–phonetic and motor knowledge of speech. We furthermore propose that nesting of cortical oscillations might be the key mechanism to understand the timing constraints that lead to the emergence of the syllable.

International audience; Knowing when an event is likely to occur allows attentional resources to be oriented toward that moment in time, enhancing processing of the event. We previously found that children (mean age 11 years) are unable to use endogenous temporal cues to orient attention in time, despite being able to use endogenous spatial cues (arrows) to orient attention in space. Arrow cues, however, may have proved beneficial by engaging exogenous (automatic), as well as endogenous (voluntary), orienting mechanisms. We therefore conducted two studies in which the exogenous properties of visual temporal cues were increased, to examine whether this helped children orient their attention in time. In the first study, the location of an imperative target was predicted by the direction of a left or right spatial arrow cue while its onset was predicted by the relative duration of a short or long temporal cue. To minimize the influence of rhythmic entrainment in the temporal condition, the foreperiod (500 ms/1100 ms) was deliberately chosen so as not to precisely match the duration of the temporal cue (100 ms/400 ms). Targets appeared either at cued locations/onset times (valid trials) or at unexpected locations/onset times (invalid trials). Adults' response times were significantly slower for invalid versus valid trials, in both spatial and temporal domains. Despite being slowed by invalid spatial cues, children (mean age 10.7 years) were unperturbed by invalid temporal cues, suggesting that these duration-based temporal cues did not help them orient attention in time. In the second study, we enhanced the exogenous properties of temporal cues further, by presenting multiple temporal cues in an isochronous (rhythmic) sequence. Again, to minimize automatic entrainment, target onset did not match the isochronous interval. Children (mean age 11.4 years), as well as adults, were now significantly slowed by invalid cues in both the temporal and spatial dimension. The sequential, as opposed to single, presentation of temporal cues therefore helped children to orient their attention in time. We suggest that the exogenous properties of sequential presentation provide a temporal scaffold that supports the additional attentional and mnemonic requirements of temporal, as compared to spatial, processing.

Action update, substituting a prepotent behavior with a new action, allows the organism to counteract surprising environmental demands. However, action update fails when the organism is uncertain about when to release the substituting behavior, when it faces temporal uncertainty. Predictive coding states that accurate perception demands minimization of precise prediction errors. Activity of the right anterior insula (rAI) is associated with temporal uncertainty. Therefore, we hypothesize that temporal uncertainty during action update would cause the anterior insula to decrease the sensitivity to ascending prediction errors. Moreover, action update requires response inhibition which recruits the frontostriatal indirect pathway associated with motor control. Therefore, we also hypothesize that temporal estimation errors modulate frontostriatal connections. To test these hypotheses, we collected fMRI data when participants performed an action-update paradigm within the context of temporal estimation. We fit dynamic causal models to the imaging data. Competing models comprised the inferior occipital gyrus, supramarginal gyrus (rSMG), rAI, right presupplementary motor area (rPreSMA), and the right striatum (rSTR). The winning model showed that temporal uncertainty drove activity into the rAI and decreased insular sensitivity to ascending prediction errors, as shown by weak connectivity strength of rSMG→rAI connections. Moreover, temporal estimation errors weakened rPreSMA→rSTR connections and also modulated rAI→rSTR connections, causing the disruption of action update. Results provide information about the neurophysiological implementation of the so-called horse-race model of action control. We suggest that, contrary to what might be believed, unsuccessful action update could be a homeostatic process that represents a Bayes optimal encoding of uncertainty.

The updating of prospective internal models is necessary to accurately predict future observations. Uncertainty-driven internal model updating has been studied using a variety of perceptual paradigms, and have revealed engagement of frontal and parietal areas. In a distinct literature, studies on temporal expectations have also characterized a time-perception network, which relies on temporal orienting of attention. However, the updating of prospective internal models is highly dependent on temporal attention, since temporal attention must be reoriented according to the current environmental demands. In this study, we used functional magnetic resonance imaging (fMRI) to evaluate to what extend the continuous manipulation of temporal prediction would recruit update-related areas and the time-perception network areas. We developed an exogenous temporal task that combines rhythm cueing and time-to-contact principles to generate implicit temporal expectation. Two patterns of motion were created: periodic (simple harmonic oscillation) and non-periodic (harmonic oscillation with variable acceleration). We found that non-periodic motion engaged the exogenous temporal orienting network, which includes the ventral premotor and inferior parietal cortices, and the cerebellum, as well as the presupplementary motor area, which has previously been implicated in internal model updating, and the motion-sensitive area MT+. Interestingly, we found a right-hemisphere preponderance suggesting the engagement of explicit timing mechanisms. We also show that the periodic motion condition, when compared to the non-periodic motion, activated a particular subset of the default-mode network (DMN) midline areas, including the left dorsomedial prefrontal cortex (DMPFC), anterior cingulate cortex (ACC), and bilateral posterior cingulate cortex/precuneus (PCC/PC). It suggests that the DMN plays a role in processing contextually expected information and supports recent evidence that the DMN may reflect the validation of prospective internal models and predictive control. Taken together, our findings suggest that continuous manipulation of temporal predictions engages representations of temporal prediction as well as task-independent updating of internal models.

Action observation is known to trigger predictions of the ongoing course of action and thus considered a hallmark example for predictive perception. A related task, which explicitly taps into the ability to predict actions based on their internal representations, is action segmentation; the task requires participants to demarcate where one action step is completed and another one begins. It thus benefits from a temporally precise prediction of the current action. Formation and exploitation of these temporal predictions of external events is now closely associated with a network including the basal ganglia and prefrontal cortex. Because decline of dopaminergic innervation leads to impaired function of the basal ganglia and prefrontal cortex in Parkinson's disease (PD), we hypothesised that PD patients would show increased temporal variability in the action segmentation task, especially under medication withdrawal (hypothesis 1). Another crucial aspect of action segmentation is its reliance on a semantic representation of actions. There is no evidence to suggest that action representations are substantially altered, or cannot be accessed, in non-demented PD patients. We therefore expected action segmentation judgments to follow the same overall patterns in PD patients and healthy controls (hypothesis 2), resulting in comparable segmentation profiles. Both hypotheses were tested with a novel classification approach. We present evidence for both hypotheses in the present study: classifier performance was slightly decreased when it was tested for its ability to predict the identity of movies segmented by PD patients, and a measure of normativity of response behaviour was decreased when patients segmented movies under medication-withdrawal without access to an episodic memory of the sequence. This pattern of results is consistent with hypothesis 1. However, the classifier analysis also revealed that responses given by patients and controls create very similar action-specific patterns, thus delivering evidence in favour hypothesis 2. In terms of methodology, the use of classifiers in the present study allowed us to establish similarity of behaviour across groups (hypothesis 2). The approach opens up a new avenue that standard statistical methods often fail to provide and is discussed in terms of its merits to measure hypothesised similarities across study populations. Kim Hannah Krause and Robert Schnuerch; German Research Foundation (DFG), Clinical Research Group (KFO)219, TP4 (GZ SCHU 1439/3-1).

Context: Establishing a neurobiological account of delusion formation that links cognitive processes, brain activity, and symptoms is important to furthering our understanding of psychosis.Objective: To explore a theoretical model of delusion formation that implicates prediction error - dependent associative learning processes in a pharmacological functional magnetic resonance imaging study using the psychotomimetic drug ketamine.Design: Within-subject, randomized, placebo-controlled study.Setting: Hospital-based clinical research facility, Addenbrooke's Hospital, Cambridge, England. The work was completed within the Wellcome Trust and Medical Research Council Behavioral and Clinical Neuroscience Institute, Cambridge.Participants: Fifteen healthy, right-handed volunteers (8 of whom were male) with a mean +/- SD age of 29 +/- 7 years and a mean +/- SD predicted full-scale IQ of 113 +/- 4 were recruited from within the local community by advertisement.Interventions: Subjects were given low-dose ketamine (100 ng/mL of plasma) or placebo while performing a causal associative learning task during functional magnetic resonance imaging. In a separate session outside the scanner, the dose was increased (to 200 ng/mL of plasma) and subjects underwent a structured clinical interview.Main Outcome Measures: Brain activation, blood plasma levels of ketamine, and scores from psychiatric ratings scales (Brief Psychiatric Ratings Scale, Present State Examination, and Clinician-Administered Dissociative States Scale).Results: Low-dose ketamine perturbs error-dependent learning activity in the right frontal cortex (P = .03). Highdose ketamine produces perceptual aberrations (P = .01) and delusion-like beliefs (P = .007). Critically, subjects showing the highest degree of frontal activation with placebo show the greatest occurrence of drug-induced perceptual aberrations (P = .03) and ideas or delusions of reference (P = .04).Conclusions: These findings relate aberrant prediction error - dependent associative learning to referential ideas and delusions via a perturbation of frontal cortical function. They are consistent with a model of delusion formation positing disruptions in error-dependent learning.

Forward predictions are not only crucial to predict somatosensory consequences of movements, but may also guide cross-modal sensory mapping (Ramnani, 2006). When a visual stimulus temporally predicts an auditory tone onset (‘when’ information), the amplitude of the N1-P2 complex is reduced compared to a temporally unpredictable tone (e.g., Ford et al., 2007; Sowman, Kuusik, & Johnson, 2012). However, cue quality and/or tone identity (‘what’ information) may further modulate visual-to- auditory predictions. When the quality of a visual cue varies, its strength to predict a subsequent tone onset influences cross- modal sensory mapping. In addition, infrequently occurring deviant tones (altered in frequency) induce uncertainty about tone identity that may further impact visual-to-auditory predictions (cf. Schwartze, Farrugia, & Kotz, 2013, for sensory predictions). A visual-to-auditory cross-modal prediction paradigm was used in the current experiment. Participants passively viewed visual cues that temporally predicted a subsequent tone onset or not. We manipulated (1) the predictive strength of a visual cue (five different cue colors vs. one [grey] color), (2) the probability of an auditory tone to occur (80% standard, 20% deviant tones), and (3) temporal predictability (predictable vs. unpredictable tone onset). Predictable blocks either preceded unpredictable blocks or vice versa (between subjects-design). This allowed investigating whether frequent standard tones are affected by uncertainty of tone identity (‘what’ information) as a function of temporal predictability (‘when’ information). The results on lateral electrodes revealed that the N1-P2 complex to standard tones was modulated by temporal predictability. The N100 amplitude was enhanced (predictable>unpredictable) when predictable blocks preceded unpredictable blocks, while the N100 amplitude was reduced (predictable