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This narrative review brings together the findings regarding the differences in the auditory event-related potentials (ERPs) between patients with depressive disorder and non-depressed control subjects. These studies' results can inform us of the possible alterations in sensory-cognitive processing in depressive disorders and the potential of using these ERPs in clinical applications. Auditory P3, mismatch negativity (MMN) and loudness dependence of auditory evoked potentials (LDAEP) were the subjects of the investigation. A search in PubMed yielded 84 studies. The findings of the reviewed studies were not highly consistent, but some patterns could be identified. For auditory P3b, the common findings were attenuated amplitude and prolonged latency among depressed patients. Regarding auditory MMN, especially the amplitude of duration deviance MMN was commonly attenuated, and the amplitude of frequency deviance MMN was increased in depressed patients. In LDAEP studies, generally, no differences between depressed patients and non-depressed controls were reported, although some group differences concerning specific depression subtypes were found. This review posits that future research should investigate whether certain stimulus conditions are particularly efficient at separating depressed and non-depressed participant groups. Future studies should contrast responses in different subpopulations of depressed patients, as well as different clinical groups (e.g., depressive disorder and anxiety disorder patients), to investigate the specificity of the auditory ERP alterations for depressive disorders. peerReviewed

Abstract. Automatic deviance detection has been widely explored in terms of mismatch responses (mismatch negativity or mismatch response) and P3a components of event-related potentials (ERPs) under a predictive coding framework; however, the somatosensory mismatch response has been investigated less often regarding the different types of changes than its auditory counterpart. It is not known whether the deviance detection responses from different modalities correlate, reflecting a general prediction error mechanism of the central nervous system. Furthermore, interoceptive functions have been associated with predictive coding theory, but whether interoceptive accuracy correlates with deviance detection brain responses has rarely been investigated. Here, we measured ERPs to changes in somatosensory stimuli’s location and intensity and in sound intensity in healthy adults ( n = 34). Interoceptive accuracy was measured with a heartbeat discrimination task, where participants indicated whether their heartbeats were simultaneous or non-simultaneous with sound stimuli. We found a mismatch response and a P3a response to somatosensory location and auditory intensity changes, but for somatosensory intensity changes, only a P3a response was found. Unexpectedly, there were neither correlations between the somatosensory location deviance and intensity deviance brain responses nor between auditory and somatosensory brain responses. In addition, the brain responses did not correlate with interoceptive accuracy. The results suggest that although deviance detection in the auditory and somatosensory modalities are likely based on similar neural mechanisms at a cellular level, their ERP indexes do not indicate a linear association in sensitivity for deviance detection between the modalities. Furthermore, although sensory deviance detection and interoceptive detection are both associated with predictive coding functions, under these experimental settings, functional relationships were not observed. These results should be taken into account in the future development of theories related to human sensory functions and in extensions of the predictive coding theory in particular.

The aim of this study was to investigate the attentional mechanism in speech processing of native and foreign language in children with and without attentional deficit. For this purpose, the P3a component, cognitive neuromarker of the attentional processes, was investigated in a two-sequence two-deviant oddball paradigm using Finnish and English speech items via event-related potentials (ERP) technique. The difference waves reflected the temporal brain dynamics of the P3a response in native and foreign language contexts. Cluster-based permutation tests evaluated the group differences over the P3a time window. A correlation analysis was conducted between the P3a response and the attention score (ATTEX) to evaluate whether the behavioral assessment reflected the neural activity. The source reconstruction method (CLARA) was used to investigate the neural origins of the attentional differences between groups and conditions. The ERP results showed a larger P3a response in the group of children with attentional problems (AP) compared to controls (CTR). The P3a response differed statistically between the two groups in the native language processing, but not in the foreign language. The ATTEX score correlated with the P3a amplitude in the native language contrasts. The correlation analyses hint at some hemispheric brain activity difference in the frontal area. The group-level CLARA reconstruction showed activation in the speech perception and attention networks over the frontal, parietal, and temporal areas. Differences in activations of these networks were found between the groups and conditions, with the AP group showing higher activity in the source level, being the origin of the ERP enhancement observed on the scalp level. peerReviewed

Funding Information: This study was supported by National Institutes of Health (Grant Nos. NIBIB RO1 EB022889, NIMH RO1 MH106174). Publisher Copyright: © 2021, The Author(s). Auditory evoked fields (AEFs) are commonly studied, yet their underlying neural mechanisms remain poorly understood. Here, we used the biophysical modelling software Human Neocortical Neurosolver (HNN) whose foundation is a canonical neocortical circuit model to interpret the cell and network mechanisms contributing to macroscale AEFs elicited by a simple tone, measured with magnetoencephalography. We found that AEFs can be reproduced by activating the neocortical circuit through a layer specific sequence of feedforward and feedback excitatory synaptic drives, similar to prior simulation of somatosensory evoked responses, supporting the notion that basic structures and activation patterns are preserved across sensory regions. We also applied the modeling framework to develop and test predictions on neural mechanisms underlying AEF differences in the left and right hemispheres, as well as in hemispheres contralateral and ipsilateral to the presentation of the auditory stimulus. We found that increasing the strength of the excitatory synaptic cortical feedback inputs to supragranular layers simulates the commonly observed right hemisphere dominance, while decreasing the input latencies and simultaneously increasing the number of cells contributing to the signal accounted for the contralateral dominance. These results provide a direct link between human data and prior animal studies and lay the foundation for future translational research examining the mechanisms underlying alteration in this fundamental biomarker of auditory processing in healthy cognition and neuropathology.

From the perspective of predictive coding, our brain embodies a hierarchical generative model to realize perception, which proactively predicts the statistical structure of sensory inputs. How are these predictive processes modified as we age? Recent research suggested that aging leads to decreased weighting of sensory inputs and increased reliance on predictions. Here we investigated whether this age-related shift from sensorium to predictions occurs at all levels of hierarchical message passing. We recorded the electroencephalography responses with an auditory local–global paradigm in a cohort of 108 healthy participants from 3 groups: seniors, adults, and adolescents. The detection of local deviancy seems largely preserved in older individuals at earlier latency (including the mismatch negativity followed by the P3a but not the reorienting negativity). In contrast, the detection of global deviancy is clearly compromised in older individuals, as they showed worse task performance and attenuated P3b. Our findings demonstrate that older brains show little decline in sensory (i.e., first-order) prediction errors but significant diminution in contextual (i.e., second-order) prediction errors. Age-related deficient maintenance of auditory information in working memory might affect whether and how lower-level prediction errors propagate to the higher level.

Speech perception is dynamic and shows changes across development. In parallel, functional differences in brain development over time have been well documented and these differences may interact with changes in speech perception during infancy and childhood. Further, there is evidence that the two hemispheres contribute unequally to speech segmentation at the sentence and phonemic levels. To disentangle those contributions, we studied the cortical tracking of various sized units of speech that are crucial for spoken language processing in children (4.7–9.3 years old, N = 34) and adults (N = 19). We measured participants’ magnetoencephalogram (MEG) responses to syllables, words, and sentences, calculated the coherence between the speech signal and MEG responses at the level of words and sentences, and further examined auditory evoked responses to syllables. Age-related differences were found for coherence values at the delta and theta frequency bands. Both frequency bands showed an effect of stimulus type, although this was attributed to the length of the stimulus and not the linguistic unit size. There was no difference between hemispheres at the source level either in coherence values for word or sentence processing or in evoked response to syllables. Results highlight the importance of the lower frequencies for speech tracking in the brain across different lexical units. Further, stimulus length affects the speech–brain associations suggesting methodological approaches should be selected carefully when studying speech envelope processing at the neural level. Speech tracking in the brain seems decoupled from more general maturation of the auditory cortex.

Predictive coding model of perception postulates that the primary objective of the brain is to infer the causes of sensory inputs by reducing prediction errors (i.e., the discrepancy between expected and actual information). Moreover, prediction errors are weighted by their precision (i.e., inverse variance), which quantifies the degree of certainty about the variables. There is accumulating evidence that the reduction of precision‐weighted prediction errors can be affected by contextual regularity (as an external factor) and selective attention (as an internal factor). However, it is unclear whether the two factors function together or separately. Here we used electroencephalography (EEG) to examine the putative interaction of contextual regularity and selective attention on this reduction process. Participants were presented with pairs of regular and irregular quartets in attended and unattended conditions. We found that contextual regularity and selective attention independently modulated the N1/MMN where the repetition effect was absent. On the P2, the two factors respectively interacted with the repetition effect without interacting with each other. The results showed that contextual regularity and selective attention likely affect the reduction of precision‐weighted prediction errors in distinct manners. While contextual regularity finetunes our efficiency at reducing precision‐weighted prediction errors, selective attention seems to modulate the reduction process following the Matthew effect of accumulated advantage. peerReviewed

Spectrotemporally complex sounds carry important information for acoustic communication. Among the important features of these sounds is the temporal duration. An event‐related potential called mismatch negativity indexes auditory change detection in humans. An analogous response (mismatch response) has been found to duration changes in speech sounds in rats but not yet in mice. We addressed whether mice show this response, and, if elicited, whether this response is functionally analogous to mismatch negativity or whether adaptation‐based models suffice to explain them. Auditory‐evoked potentials were epidurally recorded above the mice auditory cortex. The differential response to the changes in a repeated human speech sound /a/ was elicited 53–259 ms post‐change (oddball condition). The differential response was observable to the largest duration change (from 200 to 110 ms). Any smaller (from 200 to 120–180 ms at 10 ms steps) duration changes did elicit an observable response. The response to the largest duration change did not robustly differ in amplitude from the response to the change‐inducing sound presented without its repetitive background (equiprobable condition). The findings suggest that adaptation may suffice to explain responses to duration changes in spectrotemporally complex sounds in anaesthetized mice. The results pave way for development of a variety of murine models of acoustic communication. peerReviewed

According to the predictive coding model of perception, the brain constantly generates predictions of the upcoming sensory inputs. Perception is realised through a hierarchical generative model which aims at minimising the discrepancy between predictions and the incoming sensory inputs (i.e., prediction errors). Notably, prediction errors are weighted depending on precision of prior information. However, it remains unclear whether and how the brain monitors prior precision when minimising prediction errors in different contexts. The current study used magnetoencephalography (MEG) to address this question. We presented participants with repetition of two non-predicted probes embedded in context of high and low precision, namely mispredicted and unpredicted probes. Non-parametric permutation statistics showed that the minimisation of precision-weighted prediction errors started to dissociate on early components of the auditory responses (including the P1m and N1m), indicating that the brain can differentiate between these scenarios at an early stage of the auditory processing stream. Permutation statistics conducted on the depth-weighted statistical parametric maps (dSPM) source solutions of the repetition difference waves between the two non-predicted probes further revealed a cluster extending from the frontal areas to the posterior temporal areas in the left hemisphere. Overall, the results suggested that context precision not only changes the weighting of prediction errors but also modulates the dynamics of how prediction errors are minimised upon the learning of statistical regularities (achieved by stimulus repetition), which likely involves differential activation at temporal-frontal regions. peerReviewed

In humans, automatic change detection is reflected by an electrical brain response called mismatch negativity (MMN). Mismatch response is also elicited in mice, but it is unclear to what extent it is functionally similar to human MMN. We investigated this possible similarity by recording local field potentials from the auditory cortex of anesthetized mice. First, we tested whether the response to stimulus changes reflected the detection of regularity violations or adaptation to standard stimuli. Responses obtained from an oddball condition, where occasional changes in frequency were presented amongst of a standard sound, were compared to responses obtained from a control condition, where no regularities existed. To test whether the differential response to the deviant sounds in the oddball condition is dependent on sensory memory, responses from the oddball condition using 375 ms and 600 ms inter-stimulus intervals (ISI) were compared. We found a differential response to deviant sounds which was larger with the shorter than the longer ISI. Furthermore, the oddball deviant sound elicited larger response than the same sound in the control condition. These results demonstrate that the mismatch response in mice reflects detection of regularity violations and sensory memory function, as the human MMN. peerReviewed

The focus of this dissertation is to investigate the perception of single chords, both in terms of perceived emotions and psychoacoustic qualities. Previous empirical research on harmony perception has mainly been concerned with horizontal aspects of harmony: studies focusing on vertical harmony perception have been rare. The current work aims to fill some of the still evident gaps between musicology, music psychology, and psychoacoustics with regard to single chord perception by combining both empirical and theoretical approaches. The work comprises three publications, based on two large-scale empirical experiments conducted with the aim of drawing the attention of both expert and inexpert listeners to have a substantial, heterogeneous, and international participant pool. The applied chord stimuli spanned common triads and tetrads with inversions, and also more rare chord sonorities in the form of pentads and hexachords. Altogether two distinct timbres were used, piano and strings. The stimuli were analyzed in terms of psychoacoustic properties, including e.g., roughness, harmonicity, and sharpness to further account for the obtained results. The results suggest significant differences in emotion perception across different chord types. The inversions and register contributed to the evaluations significantly, non-musicians distinguishing between triadic inversions similarly to musicians. Mildly dissonant chords were more preferred than consonant chords, regardless of musical sophistication or music preferences. Certain chords, especially the major seventh chord played on the strings, conveyed the emotion of nostalgia/longing effectively. New theoretical models are suggested based on the results and psychoacoustic data to explain the capacity of certain chords to convey complex emotions, and the perception of consonance/dissonance in single chords.

Depression is associated with bias in emotional information processing, but less is known about the processing of neutral sensory stimuli. Of particular interest is processing of sound intensity which is suggested to indicate central serotonergic function. We tested weather event-related brain potentials (ERPs) to occasional changes in sound intensity can dissociate first-episode depressed, recurrent depressed and healthy control participants. The first-episode depressed showed larger N1 amplitude to deviant sounds compared to recurrent depression group and control participants. In addition, both depression groups, but not the control group, showed larger N1 amplitude to deviant than standard sounds. Whether these manifestations of sensory over-excitability in depression are directly related to the serotonergic neurotransmission requires further research. The method based on ERPs to sound intensity change is fast and low-cost way to objectively measure brain activation and holds promise as a future diagnostic tool. peerReviewed

Dyslexia, a specific reading disability, runs in families. Therefore, the risk for a child to become dyslexic increases multifold if reading difficulties occur in the family. One risk factor for dyslexia is a deficit in speech perception. Using EEG, speech sound discrimination was found to be more demanding than non- speech discrimination in typical readers in Study I. In Study II, in children with dyslexia in 3rd grade, enhanced brain responses were observed and found to be associated with better performance in reading accuracy, spelling accuracy and phonemic length discrimination tasks. The brain responses of the most accurate readers in the dyslexia group originated from a more posterior site of the right auditory cortex, suggesting the employment of compensatory brainprocessing for changes in phonemic length. In Study III, the brain responses of six-month- old infants at risk for dyslexia were found to predict their reading speed in adolescence at 14 years. The prediction was mediated via preschoolage rapid naming speed, which suggests that the same cognitive processes are needed in both fluent reading and naming. Possibly those common processes are related to automatization of the retrieval process from a mental lexicon via phonological representations. Atypical brain activation to speech in infancy seems to implicate a deficient development of phonological representations in at-risk infants that later hinders lexical access. These results indicate that even if a child is born with an increased risk for dyslexia, the risk is not insurmountable. Overall it seems that accurate speech perception, well-developed speech sound representations and automatized access to them are necessary for the development of fluent reading. This information is important in developing remediation for children at increased risk for dyslexia.