Neural Correlates of Auditory Perceptual Awareness under Informational Masking

Our ability to detect target sounds in complex acoustic backgrounds is often limited not by the ear's resolution, but by the brain's information-processing capacity. The neural mechanisms and loci of this “informational masking” are unknown. We combined magnetoencephalography with simultaneous behavioral measures in humans to investigate neural correlates of informational masking and auditory perceptual awareness in the auditory cortex. Cortical responses were sorted according to whether or not target sounds were detected by the listener in a complex, randomly varying multi-tone background known to produce informational masking. Detected target sounds elicited a prominent, long-latency response (50–250 ms), whereas undetected targets did not. In contrast, both detected and undetected targets produced equally robust auditory middle-latency, steady-state responses, presumably from the primary auditory cortex. These findings indicate that neural correlates of auditory awareness in informational masking emerge between early and late stages of processing within the auditory cortex.     

Investigating the Neural Correlates of a Streaming Percept in an Informational-Masking Paradigm

Humans routinely segregate a complex acoustic scene into different auditory streams, through the extraction of bottom-up perceptual cues and the use of top-down selective attention. To determine the neural mechanisms underlying this process, neural responses obtained through magnetoencephalography (MEG) were correlated with behavioral performance in the context of an informational masking paradigm. In half the trials, subjects were asked to detect frequency deviants in a target stream, consisting of a rhythmic tone sequence, embedded in a separate masker stream composed of a random cloud of tones. In the other half of the trials, subjects were exposed to identical stimuli but asked to perform a different task—to detect tone-length changes in the random cloud of tones. In order to verify that the normalized neural response to the target sequence served as an indicator of streaming, we correlated neural responses with behavioral performance under a variety of stimulus parameters (target tone rate, target tone frequency, and the “protection zone”, that is, the spectral area with no tones around the target frequency) and attentional states (changing task objective while maintaining the same stimuli). In all conditions that facilitated target/masker streaming behaviorally, MEG normalized neural responses also changed in a manner consistent with the behavior. Thus, attending to the target stream caused a significant increase in power and phase coherence of the responses in recording channels correlated with an increase in the behavioral performance of the listeners. Normalized neural target responses also increased as the protection zone widened and as the frequency of the target tones increased. Finally, when the target sequence rate increased, the buildup of the normalized neural responses was significantly faster, mirroring the accelerated buildup of the streaming percepts. Our data thus support close links between the perceptual and neural consequences of the auditory stream segregation.     

Neural Resolution of Formant Frequencies in the Primary Auditory Cortex of Rats

Pulse-resonance sounds play an important role in animal communication and auditory object recognition, yet very little is known about the cortical representation of this class of sounds. In this study we shine light on one simple aspect: how well does the firing rate of cortical neurons resolve resonant (“formant”) frequencies of vowel-like pulse-resonance sounds. We recorded neural responses in the primary auditory cortex (A1) of anesthetized rats to two-formant pulse-resonance sounds, and estimated their formant resolving power using a statistical kernel smoothing method which takes into account the natural variability of cortical responses. While formant-tuning functions were diverse in structure across different penetrations, most were sensitive to changes in formant frequency, with a frequency resolution comparable to that reported for rat cochlear filters.     

Audio-Visual Perception of 3D Cinematography: An fMRI Study Using Condition-Based and Computation-Based Analyses

The use of naturalistic stimuli to probe sensory functions in the human brain is gaining increasing interest. Previous imaging studies examined brain activity associated with the processing of cinematographic material using both standard “condition-based” designs, as well as “computational” methods based on the extraction of time-varying features of the stimuli (e.g. motion). Here, we exploited both approaches to investigate the neural correlates of complex visual and auditory spatial signals in cinematography. In the first experiment, the participants watched a piece of a commercial movie presented in four blocked conditions: 3D vision with surround sounds (3D-Surround), 3D with monaural sound (3D-Mono), 2D-Surround, and 2D-Mono. In the second experiment, they watched two different segments of the movie both presented continuously in 3D-Surround. The blocked presentation served for standard condition-based analyses, while all datasets were submitted to computation-based analyses. The latter assessed where activity co-varied with visual disparity signals and the complexity of auditory multi-sources signals. The blocked analyses associated 3D viewing with the activation of the dorsal and lateral occipital cortex and superior parietal lobule, while the surround sounds activated the superior and middle temporal gyri (S/MTG). The computation-based analyses revealed the effects of absolute disparity in dorsal occipital and posterior parietal cortices and of disparity gradients in the posterior middle temporal gyrus plus the inferior frontal gyrus. The complexity of the surround sounds was associated with activity in specific sub-regions of S/MTG, even after accounting for changes of sound intensity. These results demonstrate that the processing of naturalistic audio-visual signals entails an extensive set of visual and auditory areas, and that computation-based analyses can track the contribution of complex spatial aspects characterizing such life-like stimuli.     

Development of Auditory-Vocal Perceptual Skills in Songbirds

Songbirds are one of the few groups of animals that learn the sounds used for vocal communication during development. Like humans, songbirds memorize vocal sounds based on auditory experience with vocalizations of adult “tutors”, and then use auditory feedback of self-produced vocalizations to gradually match their motor output to the memory of tutor sounds. In humans, investigations of early vocal learning have focused mainly on perceptual skills of infants, whereas studies of songbirds have focused on measures of vocal production. In order to fully exploit songbirds as a model for human speech, understand the neural basis of learned vocal behavior, and investigate links between vocal perception and production, studies of songbirds must examine both behavioral measures of perception and neural measures of discrimination during development. Here we used behavioral and electrophysiological assays of the ability of songbirds to distinguish vocal calls of varying frequencies at different stages of vocal learning. The results show that neural tuning in auditory cortex mirrors behavioral improvements in the ability to make perceptual distinctions of vocal calls as birds are engaged in vocal learning. Thus, separate measures of neural discrimination and behavioral perception yielded highly similar trends during the course of vocal development. The timing of this improvement in the ability to distinguish vocal sounds correlates with our previous work showing substantial refinement of axonal connectivity in cortico-basal ganglia pathways necessary for vocal learning.     

Categorical Vowel Perception Enhances the Effectiveness and Generalization of Auditory Feedback in Human-Machine-Interfaces

Human-machine interface (HMI) designs offer the possibility of improving quality of life for patient populations as well as augmenting normal user function. Despite pragmatic benefits, utilizing auditory feedback for HMI control remains underutilized, in part due to observed limitations in effectiveness. The goal of this study was to determine the extent to which categorical speech perception could be used to improve an auditory HMI. Using surface electromyography, 24 healthy speakers of American English participated in 4 sessions to learn to control an HMI using auditory feedback (provided via vowel synthesis). Participants trained on 3 targets in sessions 1–3 and were tested on 3 novel targets in session 4. An “established categories with text cues” group of eight participants were trained and tested on auditory targets corresponding to standard American English vowels using auditory and text target cues. An “established categories without text cues” group of eight participants were trained and tested on the same targets using only auditory cuing of target vowel identity. A “new categories” group of eight participants were trained and tested on targets that corresponded to vowel-like sounds not part of American English. Analyses of user performance revealed significant effects of session and group (established categories groups and the new categories group), and a trend for an interaction between session and group. Results suggest that auditory feedback can be effectively used for HMI operation when paired with established categorical (native vowel) targets with an unambiguous cue.     

Weighting of Spatial and Spectro-Temporal Cues for Auditory Scene Analysis by Human Listeners

The auditory system creates a neuronal representation of the acoustic world based on spectral and temporal cues present at the listener''s ears, including cues that potentially signal the locations of sounds. Discrimination of concurrent sounds from multiple sources is especially challenging. The current study is part of an effort to better understand the neuronal mechanisms governing this process, which has been termed “auditory scene analysis”. In particular, we are interested in spatial release from masking by which spatial cues can segregate signals from other competing sounds, thereby overcoming the tendency of overlapping spectra and/or common temporal envelopes to fuse signals with maskers. We studied detection of pulsed tones in free-field conditions in the presence of concurrent multi-tone non-speech maskers. In “energetic” masking conditions, in which the frequencies of maskers fell within the ±1/3-octave band containing the signal, spatial release from masking at low frequencies (∼600 Hz) was found to be about 10 dB. In contrast, negligible spatial release from energetic masking was seen at high frequencies (∼4000 Hz). We observed robust spatial release from masking in broadband “informational” masking conditions, in which listeners could confuse signal with masker even though there was no spectral overlap. Substantial spatial release was observed in conditions in which the onsets of the signal and all masker components were synchronized, and spatial release was even greater under asynchronous conditions. Spatial cues limited to high frequencies (>1500 Hz), which could have included interaural level differences and the better-ear effect, produced only limited improvement in signal detection. Substantially greater improvement was seen for low-frequency sounds, for which interaural time differences are the dominant spatial cue.     

Auditory Stimuli Mimicking Ambient Sounds Drive Temporal “Delta-Brushes” in Premature Infants

In the premature infant, somatosensory and visual stimuli trigger an immature electroencephalographic (EEG) pattern, “delta-brushes,” in the corresponding sensory cortical areas. Whether auditory stimuli evoke delta-brushes in the premature auditory cortex has not been reported. Here, responses to auditory stimuli were studied in 46 premature infants without neurologic risk aged 31 to 38 postmenstrual weeks (PMW) during routine EEG recording. Stimuli consisted of either low-volume technogenic “clicks” near the background noise level of the neonatal care unit, or a human voice at conversational sound level. Stimuli were administrated pseudo-randomly during quiet and active sleep. In another protocol, the cortical response to a composite stimulus (“click” and voice) was manually triggered during EEG hypoactive periods of quiet sleep. Cortical responses were analyzed by event detection, power frequency analysis and stimulus locked averaging. Before 34 PMW, both voice and “click” stimuli evoked cortical responses with similar frequency-power topographic characteristics, namely a temporal negative slow-wave and rapid oscillations similar to spontaneous delta-brushes. Responses to composite stimuli also showed a maximal frequency-power increase in temporal areas before 35 PMW. From 34 PMW the topography of responses in quiet sleep was different for “click” and voice stimuli: responses to “clicks” became diffuse but responses to voice remained limited to temporal areas. After the age of 35 PMW auditory evoked delta-brushes progressively disappeared and were replaced by a low amplitude response in the same location. Our data show that auditory stimuli mimicking ambient sounds efficiently evoke delta-brushes in temporal areas in the premature infant before 35 PMW. Along with findings in other sensory modalities (visual and somatosensory), these findings suggest that sensory driven delta-brushes represent a ubiquitous feature of the human sensory cortex during fetal stages and provide a potential test of functional cortical maturation during fetal development.     

Selective and Efficient Neural Coding of Communication Signals Depends on Early Acoustic and Social Environment

Previous research has shown that postnatal exposure to simple, synthetic sounds can affect the sound representation in the auditory cortex as reflected by changes in the tonotopic map or other relatively simple tuning properties, such as AM tuning. However, their functional implications for neural processing in the generation of ethologically-based perception remain unexplored. Here we examined the effects of noise-rearing and social isolation on the neural processing of communication sounds such as species-specific song, in the primary auditory cortex analog of adult zebra finches. Our electrophysiological recordings reveal that neural tuning to simple frequency-based synthetic sounds is initially established in all the laminae independent of patterned acoustic experience; however, we provide the first evidence that early exposure to patterned sound statistics, such as those found in native sounds, is required for the subsequent emergence of neural selectivity for complex vocalizations and for shaping neural spiking precision in superficial and deep cortical laminae, and for creating efficient neural representations of song and a less redundant ensemble code in all the laminae. Our study also provides the first causal evidence for ‘sparse coding’, such that when the statistics of the stimuli were changed during rearing, as in noise-rearing, that the sparse or optimal representation for species-specific vocalizations disappeared. Taken together, these results imply that a layer-specific differential development of the auditory cortex requires patterned acoustic input, and a specialized and robust sensory representation of complex communication sounds in the auditory cortex requires a rich acoustic and social environment.     

On the Relevance of Natural Stimuli for the Study of Brainstem Correlates: The Example of Consonance Perception

Some combinations of musical tones sound pleasing to Western listeners, and are termed consonant, while others sound discordant, and are termed dissonant. The perceptual phenomenon of consonance has been traced to the acoustic property of harmonicity. It has been repeatedly shown that neural correlates of consonance can be found as early as the auditory brainstem as reflected in the harmonicity of the scalp-recorded frequency-following response (FFR). “Neural Pitch Salience” (NPS) measured from FFRs—essentially a time-domain equivalent of the classic pattern recognition models of pitch—has been found to correlate with behavioral judgments of consonance for synthetic stimuli. Following the idea that the auditory system has evolved to process behaviorally relevant natural sounds, and in order to test the generalizability of this finding made with synthetic tones, we recorded FFRs for consonant and dissonant intervals composed of synthetic and natural stimuli. We found that NPS correlated with behavioral judgments of consonance and dissonance for synthetic but not for naturalistic sounds. These results suggest that while some form of harmonicity can be computed from the auditory brainstem response, the general percept of consonance and dissonance is not captured by this measure. It might either be represented in the brainstem in a different code (such as place code) or arise at higher levels of the auditory pathway. Our findings further illustrate the importance of using natural sounds, as a complementary tool to fully-controlled synthetic sounds, when probing auditory perception.     

Left hemispheric dominance during auditory processing in a noisy environment

Background  

In daily life, we are exposed to different sound inputs simultaneously. During neural encoding in the auditory pathway, neural activities elicited by these different sounds interact with each other. In the present study, we investigated neural interactions elicited by masker and amplitude-modulated test stimulus in primary and non-primary human auditory cortex during ipsi-lateral and contra-lateral masking by means of magnetoencephalography (MEG).     

Effect of Mistuning on the Detection of a Tone Masked by a Harmonic Tone Complex

The human auditory system is sensitive in detecting “mistuned” components in a harmonic complex, which do not match the frequency pattern defined by the fundamental frequency of the complex. Depending on the frequency configuration, the mistuned component may be perceptually segregated from the complex and may be heard as a separate tone. In the context of a masking experiment, mistuning a single component decreases its masked threshold. In this study we propose to quantify the ability to detect a single component for fixed amounts of mistuning by adaptively varying its level. This method produces masking release by mistuning that can be compared to other masking release effects. Detection thresholds were obtained for various frequency configurations where the target component was resolved or unresolved in the auditory system. The results from 6 normal-hearing listeners show a significant decrease of masked thresholds between harmonic and mistuned conditions in all configurations and provide evidence for the employment of different detection strategies for resolved and unresolved components. The data suggest that across-frequency processing is involved in the release from masking. The results emphasize the ability of this method to assess integrative aspects of pitch and harmonicity perception.     

Role of Medio-Dorsal Frontal and Posterior Parietal Neurons during Auditory Detection Performance in Rats

To further characterize the role of frontal and parietal cortices in rat cognition, we recorded action potentials simultaneously from multiple sites in the medio-dorsal frontal cortex and posterior parietal cortex of rats while they performed a two-choice auditory detection task. We quantified neural correlates of task performance, including response movements, perception of a target tone, and the differentiation between stimuli with distinct features (different pitches or durations). A minority of units—15% in frontal cortex, 23% in parietal cortex—significantly distinguished hit trials (successful detections, response movement to the right) from correct rejection trials (correct leftward response to the absence of the target tone). Estimating the contribution of movement-related activity to these responses suggested that more than half of these units were likely signaling correct perception of the auditory target, rather than merely movement direction. In addition, we found a smaller and mostly not overlapping population of units that differentiated stimuli based on task-irrelevant details. The detection-related spiking responses we observed suggest that correlates of perception in the rat are sparsely represented among neurons in the rat''s frontal-parietal network, without being concentrated preferentially in frontal or parietal areas.     

Naturalistic Stimulus Structure Determines the Integration of Audiovisual Looming Signals in Binocular Rivalry

Rapid integration of biologically relevant information is crucial for the survival of an organism. Most prominently, humans should be biased to attend and respond to looming stimuli that signal approaching danger (e.g. predator) and hence require rapid action. This psychophysics study used binocular rivalry to investigate the perceptual advantage of looming (relative to receding) visual signals (i.e. looming bias) and how this bias can be influenced by concurrent auditory looming/receding stimuli and the statistical structure of the auditory and visual signals.Subjects were dichoptically presented with looming/receding visual stimuli that were paired with looming or receding sounds. The visual signals conformed to two different statistical structures: (1) a ‘simple’ random-dot kinematogram showing a starfield and (2) a “naturalistic” visual Shepard stimulus. Likewise, the looming/receding sound was (1) a simple amplitude- and frequency-modulated (AM-FM) tone or (2) a complex Shepard tone. Our results show that the perceptual looming bias (i.e. the increase in dominance times for looming versus receding percepts) is amplified by looming sounds, yet reduced and even converted into a receding bias by receding sounds. Moreover, the influence of looming/receding sounds on the visual looming bias depends on the statistical structure of both the visual and auditory signals. It is enhanced when audiovisual signals are Shepard stimuli.In conclusion, visual perception prioritizes processing of biologically significant looming stimuli especially when paired with looming auditory signals. Critically, these audiovisual interactions are amplified for statistically complex signals that are more naturalistic and known to engage neural processing at multiple levels of the cortical hierarchy.     

Modulatory Effects of Attention on Lateral Inhibition in the Human Auditory Cortex

Reduced neural processing of a tone is observed when it is presented after a sound whose spectral range closely frames the frequency of the tone. This observation might be explained by the mechanism of lateral inhibition (LI) due to inhibitory interneurons in the auditory system. So far, several characteristics of bottom up influences on LI have been identified, while the influence of top-down processes such as directed attention on LI has not been investigated. Hence, the study at hand aims at investigating the modulatory effects of focused attention on LI in the human auditory cortex. In the magnetoencephalograph, we present two types of masking sounds (white noise vs. withe noise passing through a notch filter centered at a specific frequency), followed by a test tone with a frequency corresponding to the center-frequency of the notch filter. Simultaneously, subjects were presented with visual input on a screen. To modulate the focus of attention, subjects were instructed to concentrate either on the auditory input or the visual stimuli. More specific, on one half of the trials, subjects were instructed to detect small deviations in loudness in the masking sounds while on the other half of the trials subjects were asked to detect target stimuli on the screen. The results revealed a reduction in neural activation due to LI, which was larger during auditory compared to visual focused attention. Attentional modulations of LI were observed in two post-N1m time intervals. These findings underline the robustness of reduced neural activation due to LI in the auditory cortex and point towards the important role of attention on the modulation of this mechanism in more evaluative processing stages.     

Interaction between Attention and Bottom-Up Saliency Mediates the Representation of Foreground and Background in an Auditory Scene

The mechanism by which a complex auditory scene is parsed into coherent objects depends on poorly understood interactions between task-driven and stimulus-driven attentional processes. We illuminate these interactions in a simultaneous behavioral–neurophysiological study in which we manipulate participants' attention to different features of an auditory scene (with a regular target embedded in an irregular background). Our experimental results reveal that attention to the target, rather than to the background, correlates with a sustained (steady-state) increase in the measured neural target representation over the entire stimulus sequence, beyond auditory attention's well-known transient effects on onset responses. This enhancement, in both power and phase coherence, occurs exclusively at the frequency of the target rhythm, and is only revealed when contrasting two attentional states that direct participants' focus to different features of the acoustic stimulus. The enhancement originates in auditory cortex and covaries with both behavioral task and the bottom-up saliency of the target. Furthermore, the target's perceptual detectability improves over time, correlating strongly, within participants, with the target representation's neural buildup. These results have substantial implications for models of foreground/background organization, supporting a role of neuronal temporal synchrony in mediating auditory object formation.     

fMRI Activity in Posterior Parietal Cortex Relates to the Perceptual Use of Binocular Disparity for Both Signal-In-Noise and Feature Difference Tasks

Visually guided action and interaction depends on the brain’s ability to (a) extract and (b) discriminate meaningful targets from complex retinal inputs. Binocular disparity is known to facilitate this process, and it is an open question how activity in different parts of the visual cortex relates to these fundamental visual abilities. Here we examined fMRI responses related to performance on two different tasks (signal-in-noise “coarse” and feature difference “fine” tasks) that have been widely used in previous work, and are believed to differentially target the visual processes of signal extraction and feature discrimination. We used multi-voxel pattern analysis to decode depth positions (near vs. far) from the fMRI activity evoked while participants were engaged in these tasks. To look for similarities between perceptual judgments and brain activity, we constructed ‘fMR-metric’ functions that described decoding performance as a function of signal magnitude. Thereafter we compared fMR-metric and psychometric functions, and report an association between judged depth and fMRI responses in the posterior parietal cortex during performance on both tasks. This highlights common stages of processing during perceptual performance on these tasks.     

Sensory-Motor Interactions for Vocal Pitch Monitoring in Non-Primary Human Auditory Cortex

The neural mechanisms underlying processing of auditory feedback during self-vocalization are poorly understood. One technique used to study the role of auditory feedback involves shifting the pitch of the feedback that a speaker receives, known as pitch-shifted feedback. We utilized a pitch shift self-vocalization and playback paradigm to investigate the underlying neural mechanisms of audio-vocal interaction. High-resolution electrocorticography (ECoG) signals were recorded directly from auditory cortex of 10 human subjects while they vocalized and received brief downward (−100 cents) pitch perturbations in their voice auditory feedback (speaking task). ECoG was also recorded when subjects passively listened to playback of their own pitch-shifted vocalizations. Feedback pitch perturbations elicited average evoked potential (AEP) and event-related band power (ERBP) responses, primarily in the high gamma (70–150 Hz) range, in focal areas of non-primary auditory cortex on superior temporal gyrus (STG). The AEPs and high gamma responses were both modulated by speaking compared with playback in a subset of STG contacts. From these contacts, a majority showed significant enhancement of high gamma power and AEP responses during speaking while the remaining contacts showed attenuated response amplitudes. The speaking-induced enhancement effect suggests that engaging the vocal motor system can modulate auditory cortical processing of self-produced sounds in such a way as to increase neural sensitivity for feedback pitch error detection. It is likely that mechanisms such as efference copies may be involved in this process, and modulation of AEP and high gamma responses imply that such modulatory effects may affect different cortical generators within distinctive functional networks that drive voice production and control.     

Two-Stage Processing of Sounds Explains Behavioral Performance Variations due to Changes in Stimulus Contrast and Selective Attention: An MEG Study

Selectively attending to task-relevant sounds whilst ignoring background noise is one of the most amazing feats performed by the human brain. Here, we studied the underlying neural mechanisms by recording magnetoencephalographic (MEG) responses of 14 healthy human subjects while they performed a near-threshold auditory discrimination task vs. a visual control task of similar difficulty. The auditory stimuli consisted of notch-filtered continuous noise masker sounds, and of 1020-Hz target tones occasionally () replacing 1000-Hz standard tones of 300-ms duration that were embedded at the center of the notches, the widths of which were parametrically varied. As a control for masker effects, tone-evoked responses were additionally recorded without masker sound. Selective attention to tones significantly increased the amplitude of the onset M100 response at 100 ms to the standard tones during presence of the masker sounds especially with notches narrower than the critical band. Further, attention modulated sustained response most clearly at 300–400 ms time range from sound onset, with narrower notches than in case of the M100, thus selectively reducing the masker-induced suppression of the tone-evoked response. Our results show evidence of a multiple-stage filtering mechanism of sensory input in the human auditory cortex: 1) one at early (100 ms) latencies bilaterally in posterior parts of the secondary auditory areas, and 2) adaptive filtering of attended sounds from task-irrelevant background masker at longer latency (300 ms) in more medial auditory cortical regions, predominantly in the left hemisphere, enhancing processing of near-threshold sounds.     

Prediction error signaling explains neuronal mismatch responses in the medial prefrontal cortex

The mismatch negativity (MMN) is a key biomarker of automatic deviance detection thought to emerge from 2 cortical sources. First, the auditory cortex (AC) encodes spectral regularities and reports frequency-specific deviances. Then, more abstract representations in the prefrontal cortex (PFC) allow to detect contextual changes of potential behavioral relevance. However, the precise location and time asynchronies between neuronal correlates underlying this frontotemporal network remain unclear and elusive. Our study presented auditory oddball paradigms along with “no-repetition” controls to record mismatch responses in neuronal spiking activity and local field potentials at the rat medial PFC. Whereas mismatch responses in the auditory system are mainly induced by stimulus-dependent effects, we found that auditory responsiveness in the PFC was driven by unpredictability, yielding context-dependent, comparatively delayed, more robust and longer-lasting mismatch responses mostly comprised of prediction error signaling activity. This characteristically different composition discarded that mismatch responses in the PFC could be simply inherited or amplified downstream from the auditory system. Conversely, it is more plausible for the PFC to exert top-down influences on the AC, since the PFC exhibited flexible and potent predictive processing, capable of suppressing redundant input more efficiently than the AC. Remarkably, the time course of the mismatch responses we observed in the spiking activity and local field potentials of the AC and the PFC combined coincided with the time course of the large-scale MMN-like signals reported in the rat brain, thereby linking the microscopic, mesoscopic, and macroscopic levels of automatic deviance detection.

Neuronal recordings in the medial prefrontal cortex of the rat demonstrate that auditory mismatch responses are purely composed of prediction error signaling activity, independent from the spectral effects that drive the auditory system.