The effects of spatial leakage correction on the reliability of EEG-based functional connectivity networks.

Electroencephalography (EEG) functional connectivity (FC) estimates are confounded by the volume conduction problem. This effect can be greatly reduced by applying FC measures insensitive to instantaneous, zero-lag dependencies (corrected measures). However, numerous studies showed that FC measures sensitive to volume conduction (uncorrected measures) exhibit higher reliability and higher subject-level identifiability. We tested how source reconstruction contributed to the reliability difference of EEG FC measures on a large (n = 201) resting-state data set testing eight FC measures (including corrected and uncorrected measures). We showed that the high reliability of uncorrected FC measures in resting state partly stems from source reconstruction: idiosyncratic noise patterns define a baseline resting-state functional network that explains a significant portion of the reliability of uncorrected FC measures. This effect remained valid for template head model-based, as well as individual head model-based source reconstruction. Based on our findings we made suggestions how to best use spatial leakage corrected and uncorrected FC measures depending on the main goals of the study.

Auditory learning of recurrent tone sequences is present in the newborn's brain.

The seemingly effortless ability of our auditory system to rapidly detect new events in a dynamic environment is crucial for survival. Whether the underlying brain processes are innate is unknown. To answer this question, electroencephalography was recorded while regularly patterned (REG) versus random (RAND) tone sequences were presented to sleeping neonates. Regular relative to random sequences elicited differential neural responses after only a single repetition of the pattern indicating the existence of an innate capacity of the auditory system to detect auditory sequential regularities. We show that the newborn auditory system accumulates evidence only somewhat longer than the minimum amount determined by the ideal Bayesian observer model (the prediction from a variable-order Markov chain model) before detecting a repeating pattern. Thus, newborns can quickly form representations for regular features of the sound input, preparing the way for learning the contingencies of the environment.

Shorter Contextual Timescale Rather Than Memory Deficit in Aging.

Many aspects of cognitive ability and brain function that change as we age look like deficits on account of measurable differences in comparison to younger adult groups. One such difference occurs in auditory sensory responses that index perceptual learning. Meta-analytic findings show reliable age-related differences in auditory responses to repetitive patterns of sound and to rare violations of those patterns, variously attributed to deficits in auditory sensory memory and inhibition. Here, we determine whether proposed deficits would render older adults less prone to primacy effects, robustly observed in young adults, which present as a tendency for first learning to have a disproportionate influence over later perceptual inference. The results confirm this reduced sensitivity to primacy effects but do not support impairment in auditory sensory memory as the origin of this difference. Instead, the aging brain produces data consistent with shorter timescales of contextual reference. In conclusion, age-related differences observed previously for perceptual inference appear highly context-specific necessitating reconsideration of whether and to what function the notion of deficit should be attributed, and even whether the notion of deficit is appropriate at all.

Predictive coding in auditory perception: challenges and unresolved questions.

Predictive coding is arguably the currently dominant theoretical framework for the study of perception. It has been employed to explain important auditory perceptual phenomena, and it has inspired theoretical, experimental and computational modelling efforts aimed at describing how the auditory system parses the complex sound input into meaningful units (auditory scene analysis). These efforts have uncovered some vital questions, addressing which could help to further specify predictive coding and clarify some of its basic assumptions. The goal of the current review is to motivate these questions and show how unresolved issues in explaining some auditory phenomena lead to general questions of the theoretical framework. We focus on experimental and computational modelling issues related to sequential grouping in auditory scene analysis (auditory pattern detection and bistable perception), as we believe that this is the research topic where predictive coding has the highest potential for advancing our understanding. In addition to specific questions, our analysis led us to identify three more general questions that require further clarification: (1) What exactly is meant by prediction in predictive coding? (2) What governs which generative models make the predictions? and (3) What (if it exists) is the correlate of perceptual experience within the predictive coding framework?

Short-term cognitive fatigue effect on auditory temporal order judgments.

Fatigue is a core symptom in many psychological disorders and it can strongly influence everyday productivity. As fatigue effects have been typically demonstrated after long hours of time on task, it was surprising that in a previous study, we accidentally found a decline of temporal order judgment (TOJ) performance within 5-8 min. After replicating prior relevant findings we tested whether pauses and/or feedback relating the participant's performance to some "standard" can eliminate or reduce this short-term performance decline. We also assessed whether the performance decline is specific to the processes evoked by the TOJ task or it is a product of either general inattentiveness or the lack of willingness to thoroughly follow the task instructions. We found that both feedback and introducing pauses between successive measurements can largely reduce the performance decline, and that these two manipulations likely mobilize overlapping capacities. Performance decline was not present in a similar task when controlling for the TOJ threshold and it was not a result of uncooperative behavior. Therefore, we conclude that the TOJ threshold decline is either specific to temporal processing in general or to the TOJ task employed in the study. Overall, the results are compatible with the notion that the decline of TOJ threshold with repeated measures represents a short-term cognitive fatigue effect. This objective fatigue measure did not correlate with subjective fatigue. The latter was rather related to perceived difficulty/effort, the reduction of positive affectivity, heightened sensitivity to criticism, and the best TOJ threshold.

Spatial cues can support auditory figure-ground segregation.

A study by Tóth, Kocsis, Háden, Szerafin, Shinn-Cunningham, and Winkler [Neuroimage 141, 108 - 119 (2016)] reported that spatial cues (such as interaural differences or ITDs) that differentiate the perceived sound source directions of a target tone sequence (figure) from simultaneous distracting tones (background) did not improve the ability of participants to detect the target sequence. The present study aims to investigate more systematically whether spatially separating a complex auditory "figure" from the background auditory stream may enhance the detection of a target in a cluttered auditory scene. Results of the presented experiment suggest that the previous negative results arose because of the specific experimental conditions tested. Here the authors find that ITDs provide a clear benefit for detecting a target tone sequence amid a mixture of other simultaneous tone bursts.

Children's perception of visual and auditory ambiguity and its link to executive functions and creativity.

The phenomenon of perceptual bistability provides insights into aspects of perceptual processing not normally accessible to everyday experience. However, most experiments have been conducted in adults, and it is not clear to what extent key aspects of perceptual switching change through development. The current research examined the ability of 6-, 8-, and 10-year-old children (N = 66) to switch between competing percepts of ambiguous visual and auditory stimuli and links between switching rate, executive functions, and creativity. The numbers of switches participants reported in two visual tasks (ambiguous figure and ambiguous structure from motion) and two auditory tasks (verbal transformation and auditory streaming) were measured in three 60-s blocks. In addition, inhibitory control was measured with a Stroop task, set shifting was measured with a verbal fluency task, and creativity was measured with a divergent thinking task. The numbers of perceptual switches increased in all four tasks from 6 to 10 years of age but differed across tasks in that they were higher in the verbal transformation and ambigous structure-from-motion tasks than in the ambigous figure and auditory streaming tasks for all age groups. Although perceptual switching rates differed across tasks, there were predictive relationships between switching rates in some tasks. However, little evidence for the influence of central processes on perceptual switching was found. Overall, the results support the notion that perceptual switching is largely modality and task specific and that this property is already evident when perceptual switching emerges.

The effects of attention and task-relevance on the processing of syntactic violations during listening to two concurrent speech streams.

The notion of automatic syntactic analysis received support from some event-related potential (ERP) studies. However, none of these studies tested syntax processing in the presence of a concurrent speech stream. Here we present two concurrent continuous speech streams, manipulating two variables potentially affecting speech processing in a fully crossed design: attention (focused vs. divided) and task (lexical - detecting numerals vs. syntactical - detecting syntactic violations). ERPs elicited by syntactic violations and numerals as targets were compared with those for distractors (task-relevant events in the unattended speech stream) and attended and unattended task-irrelevant events. As was expected, only target numerals elicited the N2b and P3 components. The amplitudes of these components did not significantly differ between focused and divided attention. Both task-relevant and task-irrelevant syntactic violations elicited the N400 ERP component within the attended but not in the unattended speech stream. P600 was only elicited by target syntactic violations. These results provide no support for the notion of automatic syntactic analysis. Rather, it appears that task-relevance is a prerequisite of P600 elicitation, implying that in-depth syntactic analysis occurs only for attended speech under everyday listening situations.

Transitional Probabilities Are Prioritized over Stimulus/Pattern Probabilities in Auditory Deviance Detection: Memory Basis for Predictive Sound Processing.

UNLABELLED: Representations encoding the probabilities of auditory events do not directly support predictive processing. In contrast, information about the probability with which a given sound follows another (transitional probability) allows predictions of upcoming sounds. We tested whether behavioral and cortical auditory deviance detection (the latter indexed by the mismatch negativity event-related potential) relies on probabilities of sound patterns or on transitional probabilities. We presented healthy adult volunteers with three types of rare tone-triplets among frequent standard triplets of high-low-high (H-L-H) or L-H-L pitch structure: proximity deviant (H-H-H/L-L-L), reversal deviant (L-H-L/H-L-H), and first-tone deviant (L-L-H/H-H-L). If deviance detection was based on pattern probability, reversal and first-tone deviants should be detected with similar latency because both differ from the standard at the first pattern position. If deviance detection was based on transitional probabilities, then reversal deviants should be the most difficult to detect because, unlike the other two deviants, they contain no low-probability pitch transitions. The data clearly showed that both behavioral and cortical auditory deviance detection uses transitional probabilities. Thus, the memory traces underlying cortical deviance detection may provide a link between stimulus probability-based change/novelty detectors operating at lower levels of the auditory system and higher auditory cognitive functions that involve predictive processing. SIGNIFICANCE STATEMENT: Our research presents the first definite evidence for the auditory system prioritizing transitional probabilities over probabilities of individual sensory events. Forming representations for transitional probabilities paves the way for predictions of upcoming sounds. Several recent theories suggest that predictive processing provides the general basis of human perception, including important auditory functions, such as auditory scene analysis. Our results demonstrate that the memory traces underlying cortical deviance detection form a link between stimulus probability-based change/novelty detectors operating at lower levels of the auditory system and higher auditory cognitive functions that involve predictive processing.

Large-scale network organization of EEG functional connectivity in newborn infants.

The organization of functional brain networks changes across human lifespan. The present study analyzed functional brain networks in healthy full-term infants (N = 139) within 1-6 days from birth by measuring neural synchrony in EEG recordings during quiet sleep. Large-scale phase synchronization was measured in six frequency bands with the Phase Lag Index. Macroscopic network organization characteristics were quantified by constructing unweighted minimum spanning tree graphs. The cortical networks in early infancy were found to be significantly more hierarchical and had a more cost-efficient organization compared with MST of random control networks, more so in the theta and alpha than in other frequency bands. Frontal and parietal sites acted as the main hubs of these networks, the topological characteristics of which were associated with gestation age (GA). This suggests that individual differences in network topology are related to cortical maturation during the prenatal period, when functional networks shift from strictly centralized toward segregated configurations. Hum Brain Mapp 38:4019-4033, 2017. © 2017 Wiley Periodicals, Inc.

EEG signatures accompanying auditory figure-ground segregation.

In everyday acoustic scenes, figure-ground segregation typically requires one to group together sound elements over both time and frequency. Electroencephalogram was recorded while listeners detected repeating tonal complexes composed of a random set of pure tones within stimuli consisting of randomly varying tonal elements. The repeating pattern was perceived as a figure over the randomly changing background. It was found that detection performance improved both as the number of pure tones making up each repeated complex (figure coherence) increased, and as the number of repeated complexes (duration) increased - i.e., detection was easier when either the spectral or temporal structure of the figure was enhanced. Figure detection was accompanied by the elicitation of the object related negativity (ORN) and the P400 event-related potentials (ERPs), which have been previously shown to be evoked by the presence of two concurrent sounds. Both ERP components had generators within and outside of auditory cortex. The amplitudes of the ORN and the P400 increased with both figure coherence and figure duration. However, only the P400 amplitude correlated with detection performance. These results suggest that 1) the ORN and P400 reflect processes involved in detecting the emergence of a new auditory object in the presence of other concurrent auditory objects; 2) the ORN corresponds to the likelihood of the presence of two or more concurrent sound objects, whereas the P400 reflects the perceptual recognition of the presence of multiple auditory objects and/or preparation for reporting the detection of a target object.

Relative Pitch Perception and the Detection of Deviant Tone Patterns.

Most people are able to recognise familiar tunes even when played in a different key. It is assumed that this depends on a general capacity for relative pitch perception; the ability to recognise the pattern of inter-note intervals that characterises the tune. However, when healthy adults are required to detect rare deviant melodic patterns in a sequence of randomly transposed standard patterns they perform close to chance. Musically experienced participants perform better than naïve participants, but even they find the task difficult, despite the fact that musical education includes training in interval recognition.To understand the source of this difficulty we designed an experiment to explore the relative influence of the size of within-pattern intervals and between-pattern transpositions on detecting deviant melodic patterns. We found that task difficulty increases when patterns contain large intervals (5-7 semitones) rather than small intervals (1-3 semitones). While task difficulty increases substantially when transpositions are introduced, the effect of transposition size (large vs small) is weaker. Increasing the range of permissible intervals to be used also makes the task more difficult. Furthermore, providing an initial exact repetition followed by subsequent transpositions does not improve performance. Although musical training correlates with task performance, we find no evidence that violations to musical intervals important in Western music (i.e. the perfect fifth or fourth) are more easily detected. In summary, relative pitch perception does not appear to be conducive to simple explanations based exclusively on invariant physical ratios.

Assessing the validity of subjective reports in the auditory streaming paradigm.

While subjective reports provide a direct measure of perception, their validity is not self-evident. Here, the authors tested three possible biasing effects on perceptual reports in the auditory streaming paradigm: errors due to imperfect understanding of the instructions, voluntary perceptual biasing, and susceptibility to implicit expectations. (1) Analysis of the responses to catch trials separately promoting each of the possible percepts allowed the authors to exclude participants who likely have not fully understood the instructions. (2) Explicit biasing instructions led to markedly different behavior than the conventional neutral-instruction condition, suggesting that listeners did not voluntarily bias their perception in a systematic way under the neutral instructions. Comparison with a random response condition further supported this conclusion. (3) No significant relationship was found between social desirability, a scale-based measure of susceptibility to implicit social expectations, and any of the perceptual measures extracted from the subjective reports. This suggests that listeners did not significantly bias their perceptual reports due to possible implicit expectations present in the experimental context. In sum, these results suggest that valid perceptual data can be obtained from subjective reports in the auditory streaming paradigm.

Newborn infants detect cues of concurrent sound segregation.

Separating concurrent sounds is fundamental for a veridical perception of one's auditory surroundings. Sound components that are harmonically related and start at the same time are usually grouped into a common perceptual object, whereas components that are not in harmonic relation or have different onset times are more likely to be perceived in terms of separate objects. Here we tested whether neonates are able to pick up the cues supporting this sound organization principle. We presented newborn infants with a series of complex tones with their harmonics in tune (creating the percept of a unitary sound object) and with manipulated variants, which gave the impression of two concurrently active sound sources. The manipulated variant had either one mistuned partial (single-cue condition) or the onset of this mistuned partial was also delayed (double-cue condition). Tuned and manipulated sounds were presented in random order with equal probabilities. Recording the neonates' electroencephalographic responses allowed us to evaluate their processing of the sounds. Results show that, in both conditions, mistuned sounds elicited a negative displacement of the event-related potential (ERP) relative to tuned sounds from 360 to 400 ms after sound onset. The mistuning-related ERP component resembles the object-related negativity (ORN) component in adults, which is associated with concurrent sound segregation. Delayed onset additionally led to a negative displacement from 160 to 200 ms, which was probably more related to the physical parameters of the sounds than to their perceptual segregation. The elicitation of an ORN-like response in newborn infants suggests that neonates possess the basic capabilities of segregating concurrent sounds by detecting inharmonic relations between the co-occurring sounds.

Processing of Horizontal Sound Localization Cues in Newborn Infants.

OBJECTIVES: By measuring event-related brain potentials (ERPs), the authors tested the sensitivity of the newborn auditory cortex to sound lateralization and to the most common cues of horizontal sound localization. DESIGN: Sixty-eight healthy full-term newborn infants were presented with auditory oddball sequences composed of frequent and rare noise segments in four experimental conditions. The authors tested in them the detection of deviations in the primary cues of sound lateralization (interaural time and level difference) and in actual sound source location (free-field and monaural sound presentation). ERP correlates of deviance detection were measured in two time windows. RESULTS: Deviations in both primary sound localization cues and the ear of stimulation elicited a significant ERP difference in the early (90 to 140 msec) time window. Deviance in actual sound source location (the free-field condition) elicited a significant response in the late (290 to 340 msec) time window. CONCLUSIONS: The early differential response may indicate the detection of a change in the respective auditory features. The authors suggest that the late differential response, which was only elicited by actual sound source location deviation, reflects the detection of location deviance integrating the various cues of sound source location. Although the results suggest that all of the tested binaural cues are processed by the neonatal auditory cortex, utilizing the cues for locating sound sources of these cues may require maturation and learning.

Modelling the emergence and dynamics of perceptual organisation in auditory streaming.

Many sound sources can only be recognised from the pattern of sounds they emit, and not from the individual sound events that make up their emission sequences. Auditory scene analysis addresses the difficult task of interpreting the sound world in terms of an unknown number of discrete sound sources (causes) with possibly overlapping signals, and therefore of associating each event with the appropriate source. There are potentially many different ways in which incoming events can be assigned to different causes, which means that the auditory system has to choose between them. This problem has been studied for many years using the auditory streaming paradigm, and recently it has become apparent that instead of making one fixed perceptual decision, given sufficient time, auditory perception switches back and forth between the alternatives-a phenomenon known as perceptual bi- or multi-stability. We propose a new model of auditory scene analysis at the core of which is a process that seeks to discover predictable patterns in the ongoing sound sequence. Representations of predictable fragments are created on the fly, and are maintained, strengthened or weakened on the basis of their predictive success, and conflict with other representations. Auditory perceptual organisation emerges spontaneously from the nature of the competition between these representations. We present detailed comparisons between the model simulations and data from an auditory streaming experiment, and show that the model accounts for many important findings, including: the emergence of, and switching between, alternative organisations; the influence of stimulus parameters on perceptual dominance, switching rate and perceptual phase durations; and the build-up of auditory streaming. The principal contribution of the model is to show that a two-stage process of pattern discovery and competition between incompatible patterns can account for both the contents (perceptual organisations) and the dynamics of human perception in auditory streaming.

What controls gain in gain control? Mismatch negativity (MMN), priors and system biases.

Repetitious patterns enable the auditory system to form prediction models specifying the most likely characteristics of subsequent sounds. Pattern deviations elicit mismatch negativity (MMN), the amplitude of which is modulated by the size of the deviation and confidence in the model. Todd et al. (Neuropsychologia 49:3399-3405, 2011; J Neurophysiol 109:99-105, 2013) demonstrated that a multi-timescale sequence reveals a bias that profoundly distorts the impact of local sound statistics on the MMN amplitude. Two sounds alternate roles as repetitious "standard" and rare "deviant" rapidly (every 0.8 min) or slowly (every 2.4 min). The bias manifests as larger MMN to the sound first encountered as deviant in slow compared to fast changing sequences, but no difference for the sound first encountered as a standard. We propose that the bias is due to how Bayesian priors shape filters of sound relevance. By examining the time-course of change in MMN amplitude we show that the bias manifests immediately after roles change but rapidly disappears thereafter. The bias was reflected in the response to deviant sounds only (not in response to standards), consistent with precision estimates extracted from second order patterns modulating gain differentially for the two sounds. Evoked responses to deviants suggest that pattern extraction and reactivation of priors can operate over tens of minutes or longer. Both MMN and deviant responses establish that: (1) priors are defined by the most proximally encountered probability distribution when one exists but; (2) when no prior exists, one is instantiated by sequence onset characteristics; and (3) priors require context interruption to be updated.

Predictive regularity representations in violation detection and auditory stream segregation: from conceptual to computational models.

Predictive accounts of perception have received increasing attention in the past 20 years. Detecting violations of auditory regularities, as reflected by the Mismatch Negativity (MMN) auditory event-related potential, is amongst the phenomena seamlessly fitting this approach. Largely based on the MMN literature, we propose a psychological conceptual framework called the Auditory Event Representation System (AERS), which is based on the assumption that auditory regularity violation detection and the formation of auditory perceptual objects are based on the same predictive regularity representations. Based on this notion, a computational model of auditory stream segregation, called CHAINS, has been developed. In CHAINS, the auditory sensory event representation of each incoming sound is considered for being the continuation of likely combinations of the preceding sounds in the sequence, thus providing alternative interpretations of the auditory input. Detecting repeating patterns allows predicting upcoming sound events, thus providing a test and potential support for the corresponding interpretation. Alternative interpretations continuously compete for perceptual dominance. In this paper, we briefly describe AERS and deduce some general constraints from this conceptual model. We then go on to illustrate how these constraints are computationally specified in CHAINS.

The effects of rhythm and melody on auditory stream segregation.

While many studies have assessed the efficacy of similarity-based cues for auditory stream segregation, much less is known about whether and how the larger-scale structure of sound sequences support stream formation and the choice of sound organization. Two experiments investigated the effects of musical melody and rhythm on the segregation of two interleaved tone sequences. The two sets of tones fully overlapped in pitch range but differed from each other in interaural time and intensity. Unbeknownst to the listener, separately, each of the interleaved sequences was created from the notes of a different song. In different experimental conditions, the notes and/or their timing could either follow those of the songs or they could be scrambled or, in case of timing, set to be isochronous. Listeners were asked to continuously report whether they heard a single coherent sequence (integrated) or two concurrent streams (segregated). Although temporal overlap between tones from the two streams proved to be the strongest cue for stream segregation, significant effects of tonality and familiarity with the songs were also observed. These results suggest that the regular temporal patterns are utilized as cues in auditory stream segregation and that long-term memory is involved in this process.