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Human observers automatically extract temporal contingencies from the environment and predict the onset of future events. Temporal predictions are modelled by the hazard function, which describes the instantaneous probability for an event to occur given it has not occurred yet. Here, we tackle the question of whether and how the human brain tracks continuous temporal hazard on a moment-to-moment basis, and how flexibly it adjusts to strictly implicit variations in the hazard function. We applied an encoding-model approach to human electroencephalographic (EEG) data recorded during a pitch-discrimination task, in which we implicitly manipulated temporal predictability of the target tones by varying the interval between cue and target tone (the foreperiod). Critically, temporal predictability was either solely driven by the passage of time (resulting in a monotonic hazard function), or was modulated to increase at intermediate foreperiods (resulting in a modulated hazard function with a peak at the intermediate foreperiod). Forward encoding models trained to predict the recorded EEG signal from different temporal hazard functions were able to distinguish between experimental conditions, showing that implicit variations of temporal hazard bear tractable signatures in the human electroencephalogram. Notably, this tracking signal was reconstructed best from the supplementary motor area (SMA), underlining this area’s link to cognitive processing of time. Our results underline the relevance of temporal hazard to cognitive processing, and show that the predictive accuracy of the encoding-model approach can be utilised to track abstract time-resolved stimuli.Significance StatementExtracting temporal predictions from sensory input allows to process future input more efficiently and to prepare responses in time. In mathematical terms, temporal predictions can be described by the hazard function, modelling the probability of an event to occur over time. Here, we show that the human EEG tracks temporal hazard in an implicit foreperiod paradigm. Forward encoding models trained to predict the recorded EEG signal from different temporal-hazard functions were able to distinguish between experimental conditions that differed in their build-up of hazard over time. These neural signatures of tracking temporal hazard converge with the extant literature on temporal processing and provide new evidence that the supplementary motor area tracks hazard under strictly implicit timing conditions.

In multi-talker situations, individuals adapt behaviorally to this listening challenge mostly with ease, but how do brain neural networks shape this adaptation? We here establish a long-sought link between large-scale neural communications in electrophysiology and behavioral success in the control of attention in difficult listening situations. In an age-varying sample of N = 154 individuals, we find that connectivity between intrinsic neural oscillations extracted from source-reconstructed electroencephalography is regulated according to the listener’s goal during a challenging dual-talker task. These dynamics occur as spatially organized modulations in power-envelope correlations of alpha and low-beta neural oscillations during approximately 2-s intervals most critical for listening behavior relative to resting-state baseline. First, left frontoparietal low-beta connectivity (16 to 24 Hz) increased during anticipation and processing of a spatial-attention cue before speech presentation. Second, posterior alpha connectivity (7 to 11 Hz) decreased during comprehension of competing speech, particularly around target-word presentation. Connectivity dynamics of these networks were predictive of individual differences in the speed and accuracy of target-word identification, respectively, but proved unconfounded by changes in neural oscillatory activity strength. Successful adaptation to a listening challenge thus latches onto two distinct yet complementary neural systems: a beta-tuned frontoparietal network enabling the flexible adaptation to attentive listening state and an alpha-tuned posterior network supporting attention to speech., This study investigates how intrinsic neural oscillations, acting in concert, tune into attentive listening. Using electroencephalography signals collected from people in a dual-talker listening task, the authors find that network connectivity of frontoparietal beta and posterior alpha oscillations is regulated according to the listener’s goal.

When listening, familiarity with an attended talker’s voice improves speech comprehension. Here, we instead investigated the effect of familiarity with a distracting talker. In an irrelevant-speech task, we assessed listeners’ working memory for the serial order of spoken digits when a task-irrelevant, distracting sentence was produced by either a familiar or an unfamiliar talker (with rare omissions of the task-irrelevant sentence). We tested two groups of listeners using the same experimental procedure. The first group were undergraduate psychology students (N=66) who had attended an introductory statistics course. Critically, each student had been taught by one of two course instructors, whose voices served as familiar and unfamiliar task-irrelevant talkers. The second group of listeners were family members and friends (N=20) who had known either one of the two talkers for more than ten years. Students, but not family members and friends, made more errors when the task-irrelevant talker was familiar versus unfamiliar. Interestingly, the effect of talker familiarity was not modulated by the presence of task-irrelevant speech: students experienced stronger working-memory disruption by a familiar talker irrespective of whether they heard a task-irrelevant sentence during memory retention or merely expected it. While previous work has shown that familiarity with an attended talker benefits speech comprehension, our findings indicate that familiarity with an ignored talker deteriorates working memory for target speech. The absence of this effect in family members and friends suggests that the degree of familiarity modulates memory disruption.

The deployment of neural alpha (8-12 Hz) lateralization in service of spatial attention is well-established: Alpha power increases in the cortical hemisphere ipsilateral to the attended hemifield, and decreases in the contralateral hemisphere, respectively. Much less is known about humans’ ability to deploy such alpha lateralization in time, and to thus exploit alpha power as a spatio-temporal filter. Here we show that spatially lateralized alpha power does signify - beyond the direction of spatial attention - the distribution of attention in time and thereby qualifies as a spatio-temporal attentional filter. Participants (N = 20) selectively listened to spoken numbers presented on one side (left vs right), while competing numbers were presented on the other side. Key to our hypothesis, temporal foreknowledge was manipulated via a visual cue, which was either instructive and indicated the to-be-probed number position (70% valid) or neutral. Temporal foreknowledge did guide participants’ attention, as they recognized numbers from the to-be-attended side more accurately following valid cues. In the magnetoencephalogram (MEG), spatial attention to the left versus right side induced lateralization of alpha power in all temporal cueing conditions. Modulation of alpha lateralization at the 0.8-Hz presentation rate of spoken numbers was stronger following instructive compared to neutral temporal cues. Critically, we found stronger modulation of lateralized alpha power specifically at the onsets of temporally cued numbers. These results suggest that the precisely timed hemispheric lateralization of alpha power qualifies as a spatio-temporal attentional filter mechanism susceptible to top-down behavioural goals.

Current models successfully describe the auditory cortical response to natural sounds with a set of spectro-temporal features. However, these models have hardly been linked to the ill-understood neurobiological changes that occur in the aging auditory cortex. Modelling the hemodynamic response to a rich natural sound mixture in N = 64 listeners of varying age, we here show that in older listeners’ auditory cortex, the key feature of temporal rate is represented with a markedly broader tuning. This loss of temporal selectivity is most prominent in primary auditory cortex and planum temporale, with no such changes in adjacent auditory or other brain areas. Amongst older listeners, we observe a direct relationship between chronological age and temporal-rate tuning, unconfounded by auditory acuity or model goodness of fit. In line with senescent neural dedifferentiation more generally, our results highlight decreased selectivity to temporal information as a hallmark of the aging auditory cortex., eLife digest It can often be difficult for an older person to understand what someone is saying, particularly in noisy environments. Exactly how and why this age-related change occurs is not clear, but it is thought that older individuals may become less able to tune in to certain features of sound. Newer tools are making it easier to study age-related changes in hearing in the brain. For example, functional magnetic resonance imaging (fMRI) can allow scientists to ‘see’ and measure how certain parts of the brain react to different features of sound. Using fMRI data, researchers can compare how younger and older people process speech. They can also track how speech processing in the brain changes with age. Now, Erb et al. show that older individuals have a harder time tuning into the rhythm of speech. In the experiments, 64 people between the ages of 18 to 78 were asked to listen to speech in a noisy setting while they underwent fMRI. The researchers then tested a computer model using the data. In the older individuals, the brain’s tuning to the timing or rhythm of speech was broader, while the younger participants were more able to finely tune into this feature of sound. The older a person was the less able their brain was to distinguish rhythms in speech, likely making it harder to understand what had been said. This hearing change likely occurs because brain cells become less specialised overtime, which can contribute to many kinds of age-related cognitive decline. This new information about why understanding speech becomes more difficult with age may help scientists develop better hearing aids that are individualised to a person’s specific needs.

An event related potential, known as the N400, has been particularly useful in investigating language processing as it serves as a neural index for semantic prediction. There are numerous studies on the functional segregation of N400 neural sources; however, the oscillatory dynamics of functional connections among the relevant sources has remained elusive. In this study we acquired magnetoencephalography data during a classic N400 paradigm, where the semantic predictability of a fixed target noun was manipulated in simple German sentences. We conducted inter‐regional functional connectivity (FC) and time–frequency analysis on known regions of the semantic network, encompassing bilateral temporal, and prefrontal cortices. Increased FC was found in less predicted (LP) nouns compared with highly predicted (HP) nouns in three connections: (a) right inferior frontal gyrus (IFG) and right middle temporal gyrus (MTG) from 0 to 300 ms mainly within the alpha band, (b) left lateral orbitofrontal (LOF) and right IFG around 400 ms within the beta band, and (c) left superior temporal gyrus (STG) and left LOF from 300 to 700 ms in the beta and low gamma bands. Furthermore, gamma spectral power (31–70 Hz) was stronger in HP nouns than in LP nouns in left anterior temporal cortices in earlier time windows (0–200 ms). Our findings support recent theories in language comprehension, suggesting fronto‐temporal top–down connections are mainly mediated through beta oscillations while gamma band frequencies are involved in matching between prediction and input.

Dopamine underlies important aspects of cognition, and has been suggested to boost cognitive performance. However, how dopamine modulates the large-scale cortical dynamics during cognitive performance has remained elusive. Using functional MRI during a working memory task in healthy young human listeners (N=22), we investigated the effect of levodopa (L-dopa) on two aspects of cortical dynamics, blood oxygen-level-dependent (BOLD) signal variability and the functional connectome of large-scale cortical networks. We here show that enhanced dopaminergic signaling modulates the two potentially interrelated aspects of large-scale cortical dynamics during cognitive performance, and the degree of these modulations is able to explain inter-individual differences in L-dopa-induced behavioral benefits. Relative to placebo, L-dopa increased BOLD signal variability in task-relevant temporal, inferior frontal, parietal and cingulate regions. On the connectome level, however, L-dopa diminished functional integration across temporal and cingulo-opercular regions. This hypo-integration was expressed as a reduction in network efficiency and modularity in more than two thirds of the participants and to different degrees. Hypo-integration co-occurred with relative hyper-connectivity in paracentral lobule and precuneus, as well as posterior putamen. Both, L-dopa-induced BOLD signal variability modulation and functional connectome modulations proved predictive of an individual’s L-dopa-induced gain in behavioral performance, namely response speed and perceptual sensitivity. Lastly, L-dopa-induced modulations of BOLD signal variability were correlated with L-dopa-induced modulation of nodal connectivity and network efficiency. Our findings underline the role of dopamine in maintaining the dynamic range of, and communication between, cortical systems, and their explanatory power for inter-individual differences in benefits from dopamine during cognitive performance.

Healthy aging is accompanied by listening difficulties, including decreased speech comprehension, that stem from an ill-understood combination of sensory and cognitive changes. Here, we use electroencephalography to demonstrate that auditory neural oscillations of older adults entrain less firmly and less flexibly to speech-paced (∼3 Hz) rhythms than younger adults’ during attentive listening. These neural entrainment effects are distinct in magnitude and origin from the neural response to sound per se. Non-entrained parieto-occipital alpha (8–12 Hz) oscillations are enhanced in young adults, but suppressed in older participants, during attentive listening. Entrained neural phase and task-induced alpha amplitude exert opposite, complementary effects on listening performance: higher alpha amplitude is associated with reduced entrainment-driven behavioural performance modulation. Thus, alpha amplitude as a task-driven, neuro-modulatory signal can counteract the behavioural corollaries of neural entrainment. Balancing these two neural strategies may present new paths for intervention in age-related listening difficulties., The changes that accompany age-related decreases in speech comprehension are not yet understood. Here, authors show that older adults are less able to entrain to speech-paced auditory rhythms and that the behavioural consequences can be counteracted by top-down neural modulation.

International audience; Can human listeners use implicit temporal contingencies in auditory input to form temporal predictions, and if so, how are these predictions represented endogenously? To assess this question, we implicitly manipulated temporal predictability in an auditory pitch discrimination task: unbeknownst to participants, the pitch of the standard tone could either be deterministically predictive of the temporal onset of the target tone, or convey no predictive information. Predictive and non-predictive conditions were presented interleaved in one stream, and separated by variable inter-stimulus intervals such that there was no dominant stimulus rhythm throughout. Even though participants were unaware of the implicit temporal contingencies, pitch discrimination sensitivity (the slope of the psychometric function) increased when the onset of the target tone was predictable in time (N = 49, 28 female, 21 male). Concurrently recorded EEG data (N = 24) revealed that standard tones that conveyed temporal predictions evoked a more negative N1 component than non-predictive standards. We observed no significant differences in oscillatory power or phase coherence between conditions during the foreperiod. Importantly, the phase angle of delta oscillations (1-3 Hz) in auditory areas in the post-standard and pre-target time windows predicted behavioral pitch discrimination sensitivity. This suggests that temporal predictions are encoded in delta oscillatory phase during the foreperiod interval. In sum, we show that auditory perception benefits from implicit temporal contingencies, and provide evidence for a role of slow neural oscillations in the endogenous representation of temporal predictions, in absence of exogenously driven entrainment to rhythmic input.

In principle, selective attention is the net result of target selection and distractor suppression. The way in which both mechanisms are implemented neurally has remained contested. Neural oscillatory power in the alpha frequency band (~10 Hz) has been implicated in the selection of to-be-attended targets, but there is lack of empirical evidence for its involvement in the suppression of to-be-ignored distractors. Here, we use electroencephalography (EEG) recordings of N = 33 human participants (males and females) to test the pre-registered hypothesis that alpha power directly relates to distractor suppression and thus operates independently from target selection. In an auditory spatial pitch discrimination task, we modulated the location (left vs right) of either a target or a distractor tone sequence, while fixing the other in the front. When the distractor was fixed in the front, alpha power relatively decreased contralaterally to the target and increased ipsilaterally. Most importantly, when the target was fixed in the front, alpha lateralization reversed in direction for the suppression of distractors on the left versus right. These data show that target-selection–independent alpha power modulation is involved in distractor suppression. While both lateralized alpha responses for selection and for suppression proved reliable, they were uncorrelated and distractor-related alpha power emerged from more anterior, frontal cortical regions. Lending functional significance to suppression-related alpha oscillations, alpha lateralization at the individual, single-trial level was predictive of behavioral accuracy. These results fuel a renewed look at neurobiological accounts of selection-independent suppressive filtering in attention.Significance statementAlthough well-established models of attention rest on the assumption that irrelevant sensory information is filtered out, the neural implementation of such a filter mechanism is unclear. Using an auditory attention task that decouples target selection from distractor suppression, we demonstrate that two sign-reversed lateralized alpha responses reflect target selection versus distractor suppression. Critically, these alpha responses are reliable, independent of each other, and generated in more anterior, frontal regions for suppression versus selection. Prediction of single-trial task performance from alpha modulation after stimulus onset agrees with the view that alpha modulation bears direct functional relevance as a neural implementation of attention. Results demonstrate that the neurobiological foundation of attention implies a selection-independent alpha oscillatory mechanism to suppress distraction.

In challenging listening conditions, closing the eyes is a strategy with intuitive appeal to improve auditory attention and perception. On the neural level, closing the eyes increases the power of alpha oscillations (∼10 Hz), which are a prime signature of auditory attention. Here, we test whether eye closure benefits neural and behavioural signatures of auditory attention and perception. Participants (N = 22) attended to one of two alternating streams of spoken numbers with open or closed eyes in a darkened chamber. After each trial, participants indicated whether probes had been among the to-be-attended or to-be-ignored numbers. In the electroencephalogram, states of relative high versus low alpha power accompanied the presentation of attended versus ignored numbers. Importantly, eye closure did not only increase the overall level of absolute alpha power but also the attentional modulation thereof. Behaviourally, however, neither perceptual sensitivity nor response criterion was affected by eye closure. To further examine whether this behavioural null-result would conceptually replicate in a simple auditory detection task, a follow-up experiment was conducted that required participants (N = 19) to detect a near-threshold target tone in noise. As in the main experiment, our results provide evidence for the absence of any difference in perceptual sensitivity and criterion for open versus closed eyes. In sum, we demonstrate here that the modulation of the human alpha rhythm by auditory attention is increased when participants close their eyes. However, our results speak against the widely held belief that eye closure per se improves listening behaviour.

Neural response adaptation plays an important role in perception and cognition. Here, we used electroencephalography to investigate how aging affects the temporal dynamics of neural adaptation in human auditory cortex. Younger (18-31 years) and older (51-70 years) normal hearing adults listened to tone sequences with varying onset-to-onset intervals. Our results show long-lasting neural adaptation such that the response to a particular tone is a nonlinear function of the extended temporal history of sound events. Most important, aging is associated with multiple changes in auditory cortex; older adults exhibit larger and less variable response magnitudes, a larger dynamic response range, and a reduced sensitivity to temporal context. Computational modeling suggests that reduced adaptation recovery times underlie these changes in the aging auditory cortex and that the extended temporal stimulation has less influence on the neural response to the current sound in older compared with younger individuals. Our human electroencephalography results critically narrow the gap to animal electrophysiology work suggesting a compensatory release from cortical inhibition accompanying hearing loss and aging.

When deciding upon a sensory stimulus, the power of prestimulus neural alpha oscillations (~10 Hz) has been shown to hold information on a perceiver's bias, or confidence, as opposed to perceptual sensitivity per se. Here, we test whether this link between prestimulus alpha power and decision confidence, previously established in vision and somatosensation, also holds in the auditory modality. Moreover, confidence usually depends on the physical evidence available in the stimulus as well as on decision accuracy. It is unclear in how far the link between prestimulus alpha power and confidence holds when physical stimulus evidence is entirely absent, and thus accuracy does not vary. We here analysed electroencephalography data from a paradigm where human listeners (N = 17) rated their confidence in the discrimination of the pitch of two tones that were, unbeknownst to the listeners, identical. Lower prestimulus alpha power as recorded at central channel sites was predictive of higher confidence ratings. Furthermore, this link was not mediated by auditory evoked activity. Our results support a direct link between prestimulus alpha power and decision confidence. This effect, first, shows up in the auditory modality similar to vision and somatosensation, and second, is present also in the complete absence of physical evidence in the stimulus and in the absence of varying accuracy. These findings speak to a model wherein low prestimulus alpha power increases neural baseline excitability, which is reflected in enhanced stimulus-evoked neural responses and higher confidence.

Increased memory load is often signified by enhanced neural oscillatory power in the alpha range (8–13 Hz), which is taken to reflect inhibition of task-irrelevant brain regions. The corresponding neural correlates of memory decay, however, are not yet well understood. In the current study, we investigated auditory short-term memory decay in humans using a delayed matching-to-sample task with pure-tone sequences. First, in a behavioral experiment, we modeled memory performance over six different delay-phase durations. Second, in a MEG experiment, we assessed alpha-power modulations over three different delay-phase durations. In both experiments, the temporal expectation for the to-be-remembered sound was manipulated so that it was either temporally expected or not. In both studies, memory performance declined over time, but this decline was weaker when the onset time of the to-be-remembered sound was expected. Similarly, patterns of alpha power in and alpha-tuned connectivity between sensory cortices changed parametrically with delay duration (i.e., decrease in occipitoparietal regions, increase in temporal regions). Temporal expectation not only counteracted alpha-power decline in heteromodal brain areas (i.e., supramarginal gyrus), but also had a beneficial effect on memory decay, counteracting memory performance decline. Correspondingly, temporal expectation also boosted alpha connectivity within attention networks known to play an active role during memory maintenance. The present data show how patterns of alpha power orchestrate short-term memory decay and encourage a more nuanced perspective on alpha power across brain space and time beyond its inhibitory role. SIGNIFICANCE STATEMENT Our sensory memories of the physical world fade quickly. We show here that this decay of short-term memory can be counteracted by so-called temporal expectation; that is, knowledge of when to expect a sensory event that an individual must remember. We also show that neural oscillations in the “alpha” (8–13 Hz) range index both the degree of memory decay (for brief sound patterns) and the respective memory benefit from temporal expectation. Spatially distributed cortical patterns of alpha power show opposing effects in auditory versus visual sensory cortices. Moreover, alpha-tuned connectivity changes within supramodal attention networks reflect the allocation of neural resources as short-term memory representations fade.

Listening requires selective neural processing of the incoming sound mixture, which in humans is borne out by a surprisingly clean representation of attended-only speech in auditory cortex. How this neural selectivity is achieved even at negative signal-to-noise ratios (SNR) remains unclear. We show that, under such conditions, a late cortical representation (i.e., neural tracking) of the ignored acoustic signal is key to successful separation of attended and distracting talkers (i.e., neural selectivity). We recorded and modelled the electroencephalographic response of 18 participants who attended to one of two simultaneously presented stories, while the SNR between the two talkers varied dynamically. The neural tracking showed an increasing early-to-late attention-biased selectivity. Importantly, acoustically dominant ignored talkers were tracked neurally by late involvement of fronto-parietal regions, which contributed to enhanced neural selectivity. This neural selectivity by way of representing the ignored talker poses a mechanistic neural account of attention under real-life acoustic conditions.

OBJECTIVES Psychoacoustic tests assessed shortly after cochlear implantation are useful predictors of the rehabilitative speech outcome. While largely independent, both spectral and temporal resolution tests are important to provide an accurate prediction of speech recognition. However, rapid tests of temporal sensitivity are currently lacking. Here, we propose a simple amplitude modulation rate discrimination (AMRD) paradigm that is validated by predicting future speech recognition in adult cochlear implant (CI) patients. DESIGN In 34 newly implanted patients, we used an adaptive AMRD paradigm, where broadband noise was modulated at the speech-relevant rate of ~4 Hz. In a longitudinal study, speech recognition in quiet was assessed using the closed-set Freiburger number test shortly after cochlear implantation (t0) as well as the open-set Freiburger monosyllabic word test 6 months later (t6). RESULTS Both AMRD thresholds at t0 (r = -0.51) and speech recognition scores at t0 (r = 0.56) predicted speech recognition scores at t6. However, AMRD and speech recognition at t0 were uncorrelated, suggesting that those measures capture partially distinct perceptual abilities. A multiple regression model predicting 6-month speech recognition outcome with deafness duration and speech recognition at t0 improved from adjusted R = 0.30 to adjusted R = 0.44 when AMRD threshold was added as a predictor. CONCLUSIONS These findings identify AMRD thresholds as a reliable, nonredundant predictor above and beyond established speech tests for CI outcome. This AMRD test could potentially be developed into a rapid clinical temporal-resolution test to be integrated into the postoperative test battery to improve the reliability of speech outcome prognosis.

Alignment of neural oscillations with temporally regular input allows listeners to generate temporal expectations. However, it remains unclear how behavior is governed in the context of temporal variability: What role do temporal expectations play, and how do they interact with the strength of neural oscillatory activity? Here, human participants detected near-threshold targets in temporally variable acoustic sequences. Temporal expectation strength was estimated using an oscillator model and pre-target neural amplitudes in auditory cortex were extracted from magnetoencephalography signals. Temporal expectations modulated target-detection performance, however, only when neural delta-band amplitudes were large. Thus, slow neural oscillations act to gate influences of temporal expectation on perception. Furthermore, slow amplitude fluctuations governed linear and quadratic influences of auditory alpha-band activity on performance. By fusing a model of temporal expectation with neural oscillatory dynamics, the current findings show that human perception in temporally variable contexts relies on complex interactions between multiple neural frequency bands.

Increased neural activity in left angular gyrus (AG) accompanies successful comprehension of acoustically degraded but highly predictable sentences, as previous functional imaging studies have shown. However, it remains unclear whether the left AG is causally relevant for the comprehension of degraded speech. Here, we applied transient virtual lesions to either the left AG or superior parietal lobe (SPL, as a control area) with repetitive transcranial magnetic stimulation (rTMS) while healthy volunteers listened to and repeated sentences with high- versus low-predictable endings and different noise vocoding levels. We expected that rTMS of AG should selectively modulate the predictability gain (i.e., the comprehension benefit from sentences with high-predictable endings) at a medium degradation level. We found that rTMS of AG indeed reduced the predictability gain at a medium degradation level of 4-band noise vocoding (relative to control rTMS of SPL). In contrast, the behavioral perturbation induced by rTMS changed with increased signal quality. Hence, at 8-band noise vocoding, rTMS over AG versus SPL decreased the number of correctly repeated keywords for sentences with low-predictable endings. Together, these results show that the degree of the rTMS interference depended jointly on signal quality and predictability. Our results provide the first causal evidence that the left AG is a critical node for facilitating speech comprehension in challenging listening conditions.

The flexible allocation of attention enables us to perceive and behave successfully despite irrelevant distractors. How do acoustic challenges influence this allocation of attention, and to what extent is this ability preserved in normally aging listeners? Younger and healthy older participants performed a masked auditory number comparison while EEG was recorded. To vary selective attention demands, we manipulated perceptual separability of spoken digits from a masking talker by varying acoustic detail (temporal fine structure). Listening conditions were adjusted individually to equalize stimulus audibility as well as the overall level of performance across participants. Accuracy increased, and response times decreased with more acoustic detail. The decrease in response times with more acoustic detail was stronger in the group of older participants. The onset of the distracting speech masker triggered a prominent contingent negative variation (CNV) in the EEG. Notably, CNV magnitude decreased parametrically with increasing acoustic detail in both age groups. Within identical levels of acoustic detail, larger CNV magnitude was associated with improved accuracy. Across age groups, neuropsychological markers further linked early CNV magnitude directly to individual attentional capacity. Results demonstrate for the first time that, in a demanding listening task, instantaneous acoustic conditions guide the allocation of attention. Second, such basic neural mechanisms of preparatory attention allocation seem preserved in healthy aging, despite impending sensory decline.

Stimulus-specific adaptation is the phenomenon whereby neural response magnitude decreases with repeated stimulation. Inconsistencies between recent nonhuman animal recordings and computational modeling suggest dynamic influences on stimulus-specific adaptation. The present human electroencephalography (EEG) study investigates the potential role of statistical context in dynamically modulating stimulus-specific adaptation by examining the auditory cortex-generated N1 and P2 components. As in previous studies of stimulus-specific adaptation, listeners were presented with oddball sequences in which the presentation of a repeated tone was infrequently interrupted by rare spectral changes taking on three different magnitudes. Critically, the statistical context varied with respect to the probability of small versus large spectral changes within oddball sequences (half of the time a small change was most probable; in the other half a large change was most probable). We observed larger N1 and P2 amplitudes (i.e., release from adaptation) for all spectral changes in the small-change compared with the large-change statistical context. The increase in response magnitude also held for responses to tones presented with high probability, indicating that statistical adaptation can overrule stimulus probability per se in its influence on neural responses. Computational modeling showed that the degree of coadaptation in auditory cortex changed depending on the statistical context, which in turn affected stimulus-specific adaptation. Thus the present data demonstrate that stimulus-specific adaptation in human auditory cortex critically depends on statistical context. Finally, the present results challenge the implicit assumption of stationarity of neural response magnitudes that governs the practice of isolating established deviant-detection responses such as the mismatch negativity.

Auditory categorization is a vital skill for perceiving the acoustic environment. Categorization depends on the discriminability of the sensory input as well as on the ability of the listener to adaptively make use of the relevant features of the sound. Previous studies on categorization have focused either on speech sounds when studying discriminability or on visual stimuli when assessing optimal cue utilization. Here, by contrast, we examined neural sensitivity to stimulus discriminability and optimal cue utilization when categorizing novel, non-speech auditory stimuli not affected by long-term familiarity. In a functional magnetic resonance imaging (fMRI) experiment, listeners categorized sounds from two category distributions, differing along two acoustic dimensions: spectral shape and duration. By introducing spectral degradation after the first half of the experiment, we manipulated both stimulus discriminability and the relative informativeness of acoustic cues. Degradation caused an overall decrease in discriminability based on spectral shape, and therefore enhanced the informativeness of duration. A relative increase in duration-cue utilization was accompanied by increased activity in left parietal cortex. Further, discriminability modulated right planum temporale activity to a higher degree when stimuli were spectrally degraded than when they were not. These findings provide support for separable contributions of parietal and posterior temporal areas to perceptual categorization. The parietal cortex seems to support the selective utilization of informative stimulus cues, while the posterior superior temporal cortex as a primarily auditory brain area supports discriminability particularly under acoustic degradation.

Background Spatial attention relatively increases the power of neural 10-Hz alpha oscillations in the hemisphere ipsilateral to attention, and decreases alpha power in the contralateral hemisphere. For gamma oscillations (>40 Hz), the opposite effect has been observed. The functional roles of lateralised oscillations for attention are currently unclear. Hypothesis If lateralised oscillations are functionally relevant for attention, transcranial stimulation of alpha versus gamma oscillations in one hemisphere should differentially modulate the accuracy of spatial attention to the ipsi-versus contralateral side. Methods 20 human participants performed a dichotic listening task under continuous transcranial alternating current stimulation (tACS, vs sham) at alpha (10 Hz) or gamma (47 Hz) frequency. On each trial, participants attended to four spoken numbers on the left or right ear, while ignoring numbers on the other ear. In order to stimulate a left temporo-parietal cortex region, which is known to show marked modulations of alpha power during auditory spatial attention, tACS (1 mA peak-to-peak amplitude) was applied at electrode positions TP7 and FC5 over the left hemisphere. Results As predicted, unihemispheric alpha-tACS relatively decreased the recall of targets contralateral to stimulation, but increased recall of ipsilateral targets. Importantly, this spatial pattern of results was reversed for gamma-tACS. Conclusions Results provide a proof of concept that transcranially stimulated oscillations can enhance spatial attention and facilitate attentional selection of speech. Furthermore, opposite effects of alpha versus gamma stimulation support the view that states of high alpha are incommensurate with active neural processing as reflected by states of high gamma.

Sensory representations, and thus human percepts, of the physical world are susceptible to fluctuations in brain state or “neural irregularity”. Furthermore, aging brains display altered levels of neural irregularity. We here show that a single, within-trial, information-theoretic measure (weighted permutation entropy) captures neural irregularity in the human electroencephalogram as a proxy for both, trait-like differences between individuals of varying age, and state-like fluctuations that bias perceptual decisions. First, the overall level of neural irregularity increased with participants’ age, paralleled by a decrease in variability over time, likely indexing age-related changes at structural and functional levels of brain activity. Second, states of higher neural irregularity were associated with optimized sensory encoding and a subsequently increased probability of choosing the first of two physically identical stimuli to be higher in pitch. In sum, neural irregularity not only characterizes behaviourally relevant brain states, but also can identify trait-like changes that come with age.

Perception of time and temporal change is critical for human cognition. Yet, perception of temporal change is susceptible to contextual influences such as changes of a sound's pitch. Using functional magnetic resonance imaging (fMRI), the current study aimed to investigate perception of temporal rate change and pitch-induced illusory distortions. In a 6 × 6 design, human participants (N=19) listened to frequency-modulated sounds (~ 4 Hz) that varied over time in both modulation rate and pitch. Participants judged the direction of rate change ('speeding up' vs. 'slowing down'), while ignoring changes in pitch. Behaviorally, rate judgments were strongly biased by pitch changes: Participants perceived rate to slow down when pitch decreased and to speed up when pitch increased ('rate-change illusion'). The fMRI data revealed activation increases with increasing task difficulty in pre-SMA, left putamen, and right IFG/insula. Importantly, activation in pre-SMA was linked to the perceptual sensitivity to discriminate rate changes and, together with the left putamen, to relative reductions in susceptibility to pitch-induced illusory distortions. Right IFG/insula activations, however, only scaled with task difficulty. These data offer a distinction between regions whose activations scale with perceptual sensitivity to features of time (pre-SMA) and those that more generally support behaving in difficult listening conditions (IFG/insula). Hence, the data underscore that individual differences in time perception can be related to different patterns of neurofunctional activation.

Slow neural oscillations (~1-15 Hz) are thought to orchestrate the neural processes of spoken language comprehension. However, functional subdivisions within this broad range of frequencies are disputed, with most studies hypothesizing only about single frequency bands. The present study utilizes an established paradigm of spoken word recognition (lexical decision) to test the hypothesis that within the slow neural oscillatory frequency range, distinct functional signatures and cortical networks can be identified at least for theta- (~3-7 Hz) and alpha-frequencies (~8-12 Hz). Listeners performed an auditory lexical decision task on a set of items that formed a word-pseudoword continuum: ranging from (1) real words over (2) ambiguous pseudowords (deviating from real words only in one vowel; comparable to natural mispronunciations in speech) to (3) pseudowords (clearly deviating from real words by randomized syllables). By means of time-frequency analysis and spatial filtering, we observed a dissociation into distinct but simultaneous patterns of alpha power suppression and theta power enhancement. Alpha exhibited a parametric suppression as items increasingly matched real words, in line with lowered functional inhibition in a left-dominant lexical processing network for more word-like input. Simultaneously, theta power in a bilateral fronto-temporal network was selectively enhanced for ambiguous pseudowords only. Thus, enhanced alpha power can neurally 'gate' lexical integration, while enhanced theta power might index functionally more specific ambiguity-resolution processes. To this end, a joint analysis of both frequency bands provides neural evidence for parallel processes in achieving spoken word recognition.

Speech signals are often compromised by disruptions originating from external (e.g., masking noise) or internal (e.g., inaccurate articulation) sources. Speech comprehension thus entails detecting and replacing missing information based on predictive and restorative neural mechanisms. The present study targets predictive mechanisms by investigating the influence of a speech segment's predictability on early, modality-specific electrophysiological responses to this segment's omission. Predictability was manipulated in simple physical terms in a single-word framework (Experiment 1) or in more complex semantic terms in a sentence framework (Experiment 2). In both experiments, final consonants of the German words Lachs ([laks], salmon) or Latz ([lats], bib) were occasionally omitted, resulting in the syllable La ([la], no semantic meaning), while brain responses were measured with multi-channel electroencephalography (EEG). In both experiments, the occasional presentation of the fragment La elicited a larger omission response when the final speech segment had been predictable. The omission response occurred ∼125–165 msec after the expected onset of the final segment and showed characteristics of the omission mismatch negativity (MMN), with generators in auditory cortical areas. Suggestive of a general auditory predictive mechanism at work, this main observation was robust against varying source of predictive information or attentional allocation, differing between the two experiments. Source localization further suggested the omission response enhancement by predictability to emerge from left superior temporal gyrus and left angular gyrus in both experiments, with additional experiment-specific contributions. These results are consistent with the existence of predictive coding mechanisms in the central auditory system, and suggestive of the general predictive properties of the auditory system to support spoken word recognition.

Transcranial magnetic stimulation (TMS) influences cortical processes. Recent findings indicate, however, that, in turn, the efficacy of TMS depends on the state of ongoing cortical oscillations. Whereas power and phase of electromyographic (EMG) activity recorded from the hand muscles as well as neural synchrony between cortex and hand muscles are known to influence the effect of TMS, to date, no study has shown an influence of the phase of cortical oscillations during wakefulness. We applied single-pulse TMS over the motor cortex and recorded motor-evoked potentials along with the electroencephalogram (EEG) and EMG. We correlated phase and power of ongoing EEG and EMG signals with the motor-evoked potential (MEP) amplitude. We also investigated the functional connectivity between cortical and hand muscle activity (corticomuscular coherence) with the MEP amplitude. EEG and EMG power and phase in a frequency band around 18 Hz correlated with the MEP amplitude. High beta-band (∼34 Hz) corticomuscular coherence exhibited a positive linear relationship with the MEP amplitude, indicating that strong synchrony between cortex and hand muscles at the moment when TMS is applied entails large MEPs. Improving upon previous studies, we demonstrate a clear dependence of TMS-induced motor effects on the state of ongoing EEG phase and power fluctuations. We conclude that not only the sampling of incoming information but also the susceptibility of cortical communication flow depends cyclically on neural phase.

Enhanced alpha power compared with a baseline can reflect states of increased cognitive load, for example, when listening to speech in noise. Can knowledge about “when” to listen (temporal expectations) potentially counteract cognitive load and concomitantly reduce alpha? The current magnetoencephalography (MEG) experiment induced cognitive load using an auditory delayed-matching-to-sample task with 2 syllables S1 and S2 presented in speech-shaped noise. Temporal expectation about the occurrence of S1 was manipulated in 3 different cue conditions: “Neutral” (uninformative about foreperiod), “early-cued” (short foreperiod), and “late-cued” (long foreperiod). Alpha power throughout the trial was highest when the cue was uninformative about the onset time of S1 (neutral) and lowest for the late-cued condition. This alpha-reducing effect of late compared with neutral cues was most evident during memory retention in noise and originated primarily in the right insula. Moreover, individual alpha effects during retention accounted best for observed individual performance differences between late-cued and neutral conditions, indicating a tradeoff between allocation of neural resources and the benefits drawn from temporal cues. Overall, the results indicate that temporal expectations can facilitate the encoding of speech in noise, and concomitantly reduce neural markers of cognitive load.

Classically, neural adaptation refers to a reduction in response magnitude by sustained stimulation. In human electroencephalography (EEG), neural adaptation has been measured, for example, as frequency-specific response decrease by previous stimulation. Only recently and mainly based on animal studies, it has been suggested that statistical properties in the stimulation lead to adjustments of neural sensitivity and affect neural response adaptation. However, it is thus far unresolved which statistical parameters in the acoustic stimulation spectrum affect frequency-specific neural adaptation, and on which time scales the effects take place. The present human EEG study investigated the potential influence of the overall spectral range as well as the spectral spacing of the acoustic stimulation spectrum on frequency-specific neural adaptation. Tones randomly varying in frequency were presented passively and computational modeling of frequency-specific neural adaptation was used. Frequency-specific adaptation was observed for all presentation conditions. Critically, however, the spread of adaptation (i.e., degree of coadaptation) in tonotopically organized regions of auditory cortex changed with the spectral range of the acoustic stimulation. In contrast, spectral spacing did not affect the spread of frequency-specific adaptation. Therefore, changes in neural sensitivity in auditory cortex are directly coupled to the overall spectral range of the acoustic stimulation, which suggests that neural adjustments to spectral stimulus statistics occur over a time scale of multiple seconds.

Neural oscillatory dynamics are a candidate mechanism to steer perception of time and temporal rate change. While oscillator models of time perception are strongly supported by behavioral evidence, a direct link to neural oscillations and oscillatory entrainment has not yet been provided. In addition, it has thus far remained unaddressed how context-induced illusory percepts of time are coded for in oscillator models of time perception. To investigate these questions, we used magnetoencephalography and examined the neural oscillatory dynamics that underpin pitch-induced illusory percepts of temporal rate change. Human participants listened to frequency-modulated sounds that varied over time in both modulation rate and pitch, and judged the direction of rate change (decrease vs increase). Our results demonstrate distinct neural mechanisms of rate perception: Modulation rate changes directly affected listeners' rate percept as well as the exact frequency of the neural oscillation. However, pitch-induced illusory rate changes were unrelated to the exact frequency of the neural responses. The rate change illusion was instead linked to changes in neural phase patterns, which allowed for single-trial decoding of percepts. That is, illusory underestimations or overestimations of perceived rate change were tightly coupled to increased intertrial phase coherence and changes in cerebro-acoustic phase lag. The results provide insight on how illusory percepts of time are coded for by neural oscillatory dynamics.

Spectral analysis of acoustic stimuli occurs in the auditory periphery (termed frequency selectivity) as well as at the level of auditory cortex (termed frequency specificity). Frequency selectivity is commonly investigated using an auditory filter model, while frequency specificity is often investigated as neural adaptation of the N1 response in electroencephalography (EEG). However, the effects of aging on frequency-specific adaptation, and the link between peripheral frequency selectivity and neural frequency specificity have not received much attention. Here, normal hearing younger (20–31 years) and older participants (49–63 years) underwent a psychophysical notched noise experiment to estimate individual auditory filters, and an EEG experiment to investigate frequency-specific adaptation in auditory cortex. The shape of auditory filters was comparable between age groups, and thus shows intact frequency selectivity in normal aging. In auditory cortex, both groups showed N1 frequency-specific neural adaptation effects that similarly varied with the spectral variance in the stimulation, while N1 responses were overall larger for older than younger participants. Importantly, the overall N1 amplitude, but not frequency-specific neural adaptation was correlated with the pass-band of the auditory filter. Thus, the current findings show a dissociation of peripheral frequency selectivity and neural frequency specificity, but suggest that widened auditory filters are compensated for by a response gain in frequency-specific areas of auditory cortex.

Under adverse listening conditions, speech comprehension profits from the expectancies that listeners derive from the semantic context. However, the neurocognitive mechanisms of this semantic benefit are unclear: How are expectancies formed from context and adjusted as a sentence unfolds over time under various degrees of acoustic degradation? In an EEG study, we modified auditory signal degradation by applying noise-vocoding (severely degraded: four-band, moderately degraded: eight-band, and clear speech). Orthogonal to that, we manipulated the extent of expectancy: strong or weak semantic context (±con) and context-based typicality of the sentence-last word (high or low: ±typ). This allowed calculation of two distinct effects of expectancy on the N400 component of the evoked potential. The sentence-final N400 effect was taken as an index of the neural effort of automatic word-into-context integration; it varied in peak amplitude and latency with signal degradation and was not reliably observed in response to severely degraded speech. Under clear speech conditions in a strong context, typical and untypical sentence completions seemed to fulfill the neural prediction, as indicated by N400 reductions. In response to moderately degraded signal quality, however, the formed expectancies appeared more specific: Only typical (+con +typ), but not the less typical (+con −typ) context–word combinations led to a decrease in the N400 amplitude. The results show that adverse listening “narrows,” rather than broadens, the expectancies about the perceived speech signal: limiting the perceptual evidence forces the neural system to rely on signal-driven expectancies, rather than more abstract expectancies, while a sentence unfolds over time.

In auditory cortex, activation and subsequent adaptation is strongest for regions responding best to a stimulated tone frequency and less for regions responding best to other frequencies. Previous attempts to characterize the spread of neural adaptation in humans investigated the auditory cortex N1 component of the event-related potentials. Importantly, however, more recent studies in animals show that neural response properties are not independent of the stimulation context. To link these findings in animals to human scalp potentials, we investigated whether contextual factors of the acoustic stimulation, namely, spectral variance, affect the spread of neural adaptation. Electroencephalograms were recorded while human participants listened to random tone sequences varying in spectral variance (narrow vs. wide). Spread of adaptation was investigated by modeling single-trial neural adaptation and subsequent recovery based on the spectro-temporal stimulation history. Frequency-specific neural responses were largest on the N1 component, and the modeled neural adaptation indices were strongly predictive of trial-by-trial amplitude variations. Yet the spread of adaption varied depending on the spectral variance in the stimulation, such that adaptation spread was broadened for tone sequences with wide spectral variance. Thus the present findings reveal context-dependent auditory cortex adaptation and point toward a flexibly adjusting auditory system that changes its response properties with the spectral requirements of the acoustic environment.

Both functional magnetic resonance imaging (fMRI) and event-related brain potential (ERP) studies have shown that verbal working memory plays an important role during sentence processing. There is growing evidence from outside of sentence processing that human alpha oscillations (7–13 Hz) play a critical role in working memory. This study aims to link this to the sentence processing domain. Time-frequency analyses and source localization were performed on electroencephalography (EEG) data that were recorded during the processing of auditorily presented sentences involving either a short or a long distance between an argument (subject or object) and the respective sentence-final verb. We reasoned that oscillatory activity in the alpha band should increase during sentences with longer argument-verb distances, since decreased temporal proximity should result in increased memory demands. When verbal working memory-intensive long-dependency sentences were compared to short-dependency sentences, a sustained oscillatory enhancement at 10 Hz was found during storage prior to the sentence-final verb, turning into a transient power increase in the beta band (13–20 Hz) at the sentence-final verb. The sources of the alpha oscillations were localized to bilaterally occipital and left parietal cortices. Only the source activity in the left parietal cortex was negatively correlated with verbal working memory abilities. These findings indicate that the parsimonious role of alpha oscillations in domain-general working memory can be extended to language, that is, sentence processing. We suggest that the function of left parietal cortex underlying verbal working memory storage during sentence processing is to inhibit the premature release of verbal information that will subsequently be integrated.

Being able to predict when an event will occur (temporal predictability) can help us prepare and time our responses. We here sought to delineate the neural and behavioural corollaries of highly implicit, probabilistic temporal predictability in an auditory foreperiod paradigm. To this end, we measured electroencephalography (EEG) and response times in two independent experiments (total N =46). Unbeknownst to participants, we induced a probabilistic variation of cue-target delays (i.e., foreperiods) in a pitch-discrimination task on a noise-embedded tone: The smaller the standard deviation of the underlying foreperiod distribution, the more predictable the time of target occurrence should be. Both experiments showed that more predictive foreperiods sped up listeners' responses. Crucially, neural signatures of temporal predictability emerged when comparing EEG activity between conditions of varying temporal predictability. First, cue-related P2 evoked responses were less pronounced for cues that implicitly signalled temporal predictability of target occurrence. Second, in both experiments, fronto-central delta (1-4Hz) phase coherence was found relatively reduced during predictive foreperiods. Concomitantly, in Experiment II, the most predictive condition yielded a central alpha (7-12Hz) power increase just before the most likely time point of tone onset, likely reflecting improved temporal orienting of attention. In sum, neural oscillations in anticipation of, and response times to a target show that humans are susceptible to even strictly implicit, probabilistic temporal regularities.

Cortical entrainment of the auditory cortex to the broadband temporal envelope of a speech signal is crucial for speech comprehension. Entrainment results in phases of high and low neural excitability, which structure and decode the incoming speech signal. Entrainment to speech is strongest in the theta frequency range (4–8 Hz), the average frequency of the speech envelope. If a speech signal is degraded, entrainment to the speech envelope is weaker and speech intelligibility declines. Besides perceptually evoked cortical entrainment, transcranial alternating current stimulation (tACS) entrains neural oscillations by applying an electric signal to the brain. Accordingly, tACS-induced entrainment in auditory cortex has been shown to improve auditory perception. The aim of the current study was to modulate speech intelligibility externally by means of tACS such that the electric current corresponds to the envelope of the presented speech stream (i.e., envelope-tACS). Participants performed the Oldenburg sentence test with sentences presented in noise in combination with envelope-tACS. Critically, tACS was induced at time lags of 0 to 250 ms in 50-ms steps relative to sentence onset (auditory stimuli were simultaneous to or preceded tACS). We performed single-subject sinusoidal, linear, and quadratic fits to the sentence comprehension performance across the time lags. We could show that the sinusoidal fit described the modulation of sentence comprehension best. Importantly, the average frequency of the sinusoidal fit was 5.12 Hz, corresponding to the peaks of the amplitude spectrum of the stimulated envelopes. This finding was supported by a significant 5-Hz peak in the average power spectrum of individual performance time series. Altogether, envelope tACS modulates intelligibility of speech in noise, presumably by enhancing and disrupting (time lag with in-or out-of-phase stimulation, respectively) cortical entrainment to the speech envelope in auditory cortex.

ObjectiveConventional, multi-channel scalp electroencephalography (EEG) allows the identification of the attended speaker in concurrent-listening (“cocktail party”) scenarios. This implies that EEG might provide valuable information to complement hearing aids with some form of EEG and to install a level of neuro-feedback.ApproachTo investigate whether a listener’s attentional focus can be detected from single-channel hearing-aid-compatible EEG configurations, we recorded EEG from three electrodes inside the ear canal (“in-Ear-EEG”) and additionally from 64 electrodes on the scalp. In two different, concurrent listening tasks, participants (n = 7) were fitted with individualized in-Ear-EEG pieces and were either asked to attend to one of two dichotically-presented, concurrent tone streams or to one of two diotically-presented, concurrent audiobooks. A forward encoding model was trained to predict the EEG response at single EEG channels.Main resultsEach individual participants’ attentional focus could be detected from single-channel EEG response recorded from short-distance configurations consisting only of a single in-Ear-EEG electrode and an adjacent scalp-EEG electrode. The differences in neural responses to attended and ignored stimuli were consistent in morphology (i.e., polarity and latency of components) across subjects.SignificanceIn sum, our findings show that the EEG response from a single-channel, hearing-aid-compatible configuration provides valuable information to identify a listener’s focus of attention.

Contains fulltext : 122173.pdf (Publisher’s version ) (Open Access) Listeners show a remarkable ability to quickly adjust to degraded speech input. Here, we aimed to identify the neural mechanisms of such short-term perceptual adaptation. In a sparse-sampling, cardiac-gated functional magnetic resonance imaging (fMRI) acquisition, human listeners heard and repeated back 4-band-vocoded sentences (in which the temporal envelope of the acoustic signal is preserved, while spectral information is highly degraded). Clear-speech trials were included as baseline. An additional fMRI experiment on amplitude modulation rate discrimination quantified the convergence of neural mechanisms that subserve coping with challenging listening conditions for speech and non-speech. First, the degraded speech task revealed an "executive" network (comprising the anterior insula and anterior cingulate cortex), parts of which were also activated in the non-speech discrimination task. Second, trial-by-trial fluctuations in successful comprehension of degraded speech drove hemodynamic signal change in classic "language" areas (bilateral temporal cortices). Third, as listeners perceptually adapted to degraded speech, downregulation in a cortico-striato-thalamo-cortical circuit was observable. The present data highlight differential upregulation and downregulation in auditory-language and executive networks, respectively, with important subcortical contributions when successfully adapting to a challenging listening situation. 10 p.

Advances in brain-computer interface research have recently empowered the development of wearable sensors to record mobile electroencephalography (EEG) as an unobtrusive and easy-to-use alternative to conventional scalp EEG. One such mobile solution is to record EEG from the ear canal, which has been validated for auditory steady state responses and discrete event related potentials (ERPs). However, it is still under discussion where to place recording and reference electrodes to capture best responses to auditory stimuli. Furthermore, the technology has not yet been tested and validated for ecologically relevant auditory stimuli such as speech. In this study, Ear-EEG and conventional scalp EEG were recorded simultaneously in a discrete-tone as well as a continuous-speech design. The discrete stimuli were applied in a dichotic oddball paradigm, while continuous stimuli were presented diotically as two simultaneous talkers. Cross-correlation of stimulus envelope and Ear-EEG was assessed as a measure of ongoing neural tracking. The extracted ERPs from Ear-EEG revealed typical auditory components yet depended critically on the reference electrode chosen. Reliable neural-tracking responses were extracted from the Ear-EEG for both paradigms, albeit weaker in amplitude than from scalp EEG. In conclusion, this study shows the feasibility of extracting relevant neural features from ear-canal-recorded "Ear-EEG", which might augment future hearing technology.

A number of findings indicate gender differences in language-related functional hemispheric brain asymmetry. To test if such gender-specific laterality is already present at the level of vowel-processing, the auditory evoked magnetic field was recorded in healthy right-handed male and female participants in response to the German synthetic vowels [a], [e] and [i]. Female participants exhibited stronger N100m responses than male participants over the left hemisphere. This observation was highly reliable across repeated experimental sessions. The present lateralization shows that previous findings suggesting a stronger left-hemispheric dominance for verbal material in males than in females can not be generalized to basic speech elements. Furthermore, the present results support the importance of controlling for gender ratio in studies of phonetic processing.

Speech comprehension is a complex human skill, the performance of which requires the perceiver to combine information from several sources - e.g. voice, face, gesture, linguistic context - to achieve an intelligible and interpretable percept. We describe a functional imaging investigation of how auditory, visual and linguistic information interact to facilitate comprehension. Our specific aims were to investigate the neural responses to these different information sources, alone and in interaction, and further to use behavioural speech comprehension scores to address sites of intelligibility-related activation in multifactorial speech comprehension. In fMRI, participants passively watched videos of spoken sentences, in which we varied Auditory Clarity (with noise-vocoding), Visual Clarity (with Gaussian blurring) and Linguistic Predictability. Main effects of enhanced signal with increased auditory and visual clarity were observed in overlapping regions of posterior STS. Two-way interactions of the factors (auditory × visual, auditory × predictability) in the neural data were observed outside temporal cortex, where positive signal change in response to clearer facial information and greater semantic predictability was greatest at intermediate levels of auditory clarity. Overall changes in stimulus intelligibility by condition (as determined using an independent behavioural experiment) were reflected in the neural data by increased activation predominantly in bilateral dorsolateral temporal cortex, as well as inferior frontal cortex and left fusiform gyrus. Specific investigation of intelligibility changes at intermediate auditory clarity revealed a set of regions, including posterior STS and fusiform gyrus, showing enhanced responses to both visual and linguistic information. Finally, an individual differences analysis showed that greater comprehension performance in the scanning participants (measured in a post-scan behavioural test) were associated with increased activation in left inferior frontal gyrus and left posterior STS. The current multimodal speech comprehension paradigm demonstrates recruitment of a wide comprehension network in the brain, in which posterior STS and fusiform gyrus form sites for convergence of auditory, visual and linguistic information, while left-dominant sites in temporal and frontal cortex support successful comprehension.

In language processing, the relative contribution of early sensory and higher cognitive brain areas is still an open issue. A recent controversial hypothesis proposes that sensory cortices show sensitivity to syntactic processes, whereas other studies suggest a wider neural network outside sensory regions. The goal of the current event‐related fMRI study is to clarify the contribution of sensory cortices in auditory syntactic processing in a 2 × 2 design. Two‐word utterances were presented auditorily and varied both in perceptual markedness (presence or absence of an overt word category marking “‐t”), and in grammaticality (syntactically correct or incorrect). A multivariate pattern classification approach was applied to the data, flanked by conventional cognitive subtraction analyses. The combination of methods and the 2 × 2 design revealed a clear picture: The cognitive subtraction analysis found initial syntactic processing signatures in a neural network including the left IFG, the left aSTG, the left superior temporal sulcus (STS), as well as the right STS/STG. Classification of local multivariate patterns indicated the left‐hemispheric regions in IFG, aSTG, and STS to be more syntax‐specific than the right‐hemispheric regions. Importantly, auditory sensory cortices were only sensitive to the overt perceptual marking, but not to the grammaticality, speaking against syntax‐inflicted sensory cortex modulations. Instead, our data provide clear evidence for a distinction between regions involved in pure perceptual processes and regions involved in initial syntactic processes. Hum Brain Mapp, 2012. © 2011 Wiley Periodicals, Inc.

When listening to speech under adverse conditions, expectancies resulting from semantic context can have a strong impact on comprehension. Here we ask how minimal variations in semantic context (cloze probability) affect the unfolding comprehension of acoustically degraded speech. Three main results are observed in the brain electric response. First, auditory evoked responses to a degraded sentence's onset (N100) correlate with participants' comprehension scores, but are generally more vigorous for more degraded sentences. Second, a pronounced N400 in response to low-cloze sentence-final words, reflecting the integration effort of words into context, increases linearly with improving speech intelligibility. Conversely, transient enhancement in Gamma band power (γ, ~40-70 Hz) during high-cloze sentence-final words (~600 ms) reflects top-down-facilitated integration. This γ-band effect also varies parametrically with signal quality. Third, a negative correlation of N100 amplitude at sentence onset and the later γ-band response is found in moderately degraded speech. This reflects two partly distinct neural strategies when dealing with moderately degraded speech; a more "bottom-up," resource-allocating, and effortful versus a more "top-down," associative and facilitatory strategy. Results also emphasize the non-redundant contributions of phase-locked (evoked) and non-phase-locked (induced) oscillatory brain dynamics in auditory EEG.

In speech comprehension, the processing of auditory information and linguistic context are mutually dependent. This functional magnetic resonance imaging study examines how semantic expectancy ("cloze probability") in variably intelligible sentences ("noise vocoding") modulates the brain bases of comprehension. First, intelligibility-modulated activation along the superior temporal sulci (STS) was extended anteriorly and posteriorly in low-cloze sentences (e.g., "she weighs the flour") but restricted to a mid-superior temporal gyrus/STS area in more predictable high-cloze sentences (e.g., "she sifts the flour"). Second, the degree of left inferior frontal gyrus (IFG) (Brodmann's area 44) involvement in processing low-cloze constructions was proportional to increasing intelligibility. Left inferior parietal cortex (IPC; angular gyrus) activation accompanied successful speech comprehension that derived either from increased signal quality or from semantic facilitation. The results show that successful decoding of speech in auditory cortex areas regulates language-specific computation (left IFG and IPC). In return, semantic expectancy can constrain these speech-decoding processes, with fewer neural resources being allocated to highly predictable sentences. These findings offer an important contribution toward the understanding of the functional neuroanatomy in speech comprehension.

Psychophysical target detection has been shown to be modulated by slow oscillatory brain phase. However, thus far, only low-level sensory stimuli have been used as targets. The current human electroencephalography (EEG) study examined the influence of neural oscillatory phase on a lexical-decision task performed for stimuli embedded in noise. Neural phase angles were compared for correct versus incorrect lexical decisions using a phase bifurcation index (BI), which quantifies differences in mean phase angles and phase concentrations between correct and incorrect trials. Neural phase angles in the alpha frequency range (8–12 Hz) over right anterior sensors were approximately antiphase in a prestimulus time window, and thus successfully distinguished between correct and incorrect lexical decisions. Moreover, alpha-band oscillations were again approximately antiphase across participants for correct versus incorrect trials during a later peristimulus time window (∼500 ms) at left-central electrodes. Strikingly, lexical decision accuracy was not predicted by either event-related potentials (ERPs) or oscillatory power measures. We suggest that correct lexical decisions depend both on successful sensory processing, which is made possible by the alignment of stimulus onset with an optimal alpha phase, as well as integration and weighting of decisional information, which is coupled to alpha phase immediately following the critical manipulation that differentiated words from pseudowords. The current study constitutes a first step toward characterizing the role of dynamic oscillatory brain states for higher cognitive functions, such as spoken word recognition.

We investigated the functional neuroanatomy of vowel processing. We compared attentive auditory perception of natural German vowels to perception of nonspeech band‐passed noise stimuli using functional magnetic resonance imaging (fMRI). More specifically, the mapping in auditory cortex of first and second formants was considered, which spectrally characterize vowels and are linked closely to phonological features. Multiple exemplars of natural German vowels were presented in sequences alternating either mainly along the first formant (e.g., [u]‐[o], [i]‐[e]) or along the second formant (e.g., [u]‐[i], [o]‐[e]). In fixed‐effects and random‐effects analyses, vowel sequences elicited more activation than did nonspeech noise in the anterior superior temporal cortex (aST) bilaterally. Partial segregation of different vowel categories was observed within the activated regions, suggestive of a speech sound mapping across the cortical surface. Our results add to the growing evidence that speech sounds, as one of the behaviorally most relevant classes of auditory objects, are analyzed and categorized in aST. These findings also support the notion of an auditory “what” stream, with highly object‐specialized areas anterior to primary auditory cortex. Hum. Brain Mapp, 2005. © 2005 Wiley‐Liss, Inc.

The present study used magnetic source imaging to examine gender differences in the functional hemispheric asymmetry of auditory processing. The auditory evoked N100m was examined in male and female subjects in response to natural syllables with varying consonant and vowel as well as nonspeech noise. In an additional task subjects had to categorize different syllables from the first 35 ms of syllables, that is, the plosive and the formant transition. Syllable-evoked N100m activity was larger in the left than in the right hemisphere in female but not in male subjects. This gender-specific hemispheric asymmetry was speech specific, that is, absent when processing meaningless noise. Only in females did the degree of left-lateralization predict successful syllable categorization from short syllable bursts: Results suggest gender-specific differences in spectro-temporal analysis of speech.

The present study examined the cortical activity during processing of vocalic segments by means of whole-head magnetoencephalography (MEG) to see whether respective cortical maps are stable across repeated measurements. We investigated the spatial configuration and temporal characteristics of the N100m generators of the auditory-evoked field during the processing of the synthetic German vowels [a], [e] and [i] across 10 repeated measurements in a single subject. Between vowels, N100m latency as well as source location differences were found with the latency differences being in accordance with tonochronic principles. The spatial configuration of the different vowel sources was related to differences in acoustic/phonological features. Vowels differing maximally in those features, i.e., [a] and [i], showed larger Euclidean distances between N100m vowel sources than [e] and [i]. This pattern was repeatable across sessions and independent of the source modeling strategy for left-hemispheric data. Compared to a pure tone control condition, the N100m generators of vowels were localized in more anterior, superior and lateral parts of the temporal lobe and showed longer latencies. Being aware of the limited significance of conclusions drawn from a single case study, the study yielded a repeatable spatial and temporal pattern of vowel source activity in the auditory cortex which was determined by the distinctiveness of the formant frequencies corresponding to abstract phonological features.

This study further elucidates determinants of vowel perception in the human auditory cortex. The vowel inventory of a given language can be classified on the basis of phonological features which are closely linked to acoustic properties. A cortical representation of speech sounds based on these phonological features might explain the surprisingly inverse correlation between immense variance in the acoustic signal and high accuracy of speech recognition. We investigated timing and mapping of the N100m elicited by 42 tokens of seven natural German vowels varying along the phonological features tongue height (corresponding to the frequency of the first formant) and place of articulation (corresponding to the frequency of the second and third formants). Auditoryevoked fields were recorded using a 148-channel whole-head magnetometer while subjects performed target vowel detection tasks. Source location differences appeared to be driven by place of articulation: Vowels with mutually exclusive place of articulation features, namely, coronal and dorsal elicited separate centers of activation along the posterior-anterior axis. Additionally, the time course of activation as reflected in the N100m peak latency distinguished between vowel categories especially when the spatial distinctiveness of cortical activation was low. In sum, results suggest that both N100m latency and source location as well as their interaction reflect properties of speech stimuli that correspond to abstract phonological features.

Temporal expectations enhance neural encoding precision, reflected in optimized alignment of slow neural oscillatory phase, and facilitate subsequent stimulus processing. If an event's exact occurrence time is unknown, temporal expectations arise solely from the passage of time. Here, we show that this specific type of temporal expectation is also reflected in neural phase organization. While undergoing magnetoencephalography, participants performed an auditory-delayed matching-to-sample task with two syllables (S1, S2). Critically, S1-onset time varied in the 0.6-1.8-s (i.e., 0.6-1.7 Hz) range. Increasing S1-onset times led to increased slow-delta (0.6-0.9 Hz) phase coherence over right frontotemporal sensors during S1 encoding. Moreover, individuals with higher slow-delta coherence showed decreased alpha power (8-13 Hz) during subsequent memory retention. In sum, temporal expectations based on the passage of time optimize the precise alignment of neural oscillatory phase with an expected stimulus.