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Time implicitly shapes cognition, but time is also explicitly represented, for instance, in the form of durations. Parsimoniously, the brain could use the same mechanisms for implicit and explicit timing. Yet, the evidence has been equivocal, revealing both joint versus separate signatures of timing. Here, we directly compared implicit and explicit timing using magnetoencephalography, whose temporal resolution allows investigating the different stages of the timing processes. Implicit temporal predictability was induced in an auditory paradigm by a manipulation of the foreperiod. Participants received two consecutive task instructions: discriminate pitch (indirect measure of implicit timing) or duration (direct measure of explicit timing). The results show that the human brain efficiently extracts implicit temporal statistics of sensory environments, to enhance the behavioral and neural responses to auditory stimuli, but that those temporal predictions did not improve explicit timing. In both tasks, attentional orienting in time during predictive foreperiods was indexed by an increase in alpha power over visual and parietal areas. Furthermore, pretarget induced beta power in sensorimotor and parietal areas increased during implicit compared to explicit timing, in line with the suggested role for beta oscillations in temporal prediction. Interestingly, no distinct neural dynamics emerged when participants explicitly paid attention to time, compared to implicit timing. Our work thus indicates that implicit timing shapes the behavioral and sensory response in an automatic way and is reflected in oscillatory neural dynamics, whereas the translation of implicit temporal statistics to explicit durations remains somewhat inconclusive, possibly because of the more abstract nature of this task.

Listening in everyday life requires attention to be deployed dynamically – when listening is expected to be difficult and when relevant information is expected to occur – to conserve mental resources. Conserving mental resources may be particularly important for older adults who often experience difficulties understanding speech. In the current study, we use electro- and magnetoencephalography to investigate the neural and behavioral mechanics of attention regulation during listening and the effects that aging has on these. We first show in younger adults (17–31 years) that neural alpha oscillatory activity indicates when in time attention is deployed (Experiment 1) and that deployment depends on listening difficulty (Experiment 2). Experiment 3 investigated age-related changes in auditory attention regulation. Middle-aged and older adults (54–72 years) show successful attention regulation but appear to utilize timing information differently compared to younger adults (20–33 years). We show a notable age-group dissociation in recruited brain regions. In younger adults, superior parietal cortex underlies alpha power during attention regulation, whereas, in middle-aged and older adults, alpha power emerges from more ventro-lateral areas (posterior temporal cortex). This difference in the sources of alpha activity between age groups only occurred during task performance and was absent during rest (Experiment S1). In sum, our study suggests that middle-aged and older adults employ different neural control strategies compared to younger adults to regulate attention in time under listening challenges.

Speech comprehension is often challenged by increased background noise, but can be facilitated via the semantic context of a sentence. This predictability gain relies on an interplay of language-specific semantic and domain-general brain regions. However, age-related differences in the interactions within and between semantic and domain-general networks remain poorly understood. Using functional neuroimaging, we investigated commonalities and differences in network interactions enabling processing of degraded speech in healthy young and old participants. Participants performed a sentence repetition task while listening to sentences with high and low predictable endings and varying intelligibility. Stimulus intelligibility was adjusted to individual hearing abilities. Older adults showed an undiminished behavioural predictability gain. Likewise, both groups recruited a similar set of semantic and cingulo-opercular brain regions. However, we observed age-related differences in effective connectivity for high predictable speech of increasing intelligibility. Young adults exhibited stronger connectivity between regions of the cingulo-opercular network and between left insula and the posterior middle temporal gyrus. Moreover, these interactions were excitatory in young adults but inhibitory in old adults. Finally, the degree of the inhibitory influence between cingulo-opercular regions was predictive of the behavioural sensitivity towards changes in intelligibility for high predictable sentences in older adults only. Our results demonstrate that the predictability gain is relatively preserved in older adults when stimulus intelligibility is individually adjusted. While young and old participants recruit similar brain regions, differences manifest in underlying network interactions. Together, these results suggest that ageing affects the network configuration rather than regional activity during successful speech comprehension under challenging listening conditions.

Humans are born into a social environment and from early on possess a range of abilities to detect and respond to social cues. In the past decade, there has been a rapidly increasing interest in investigating the neural responses underlying such early social processes under naturalistic conditions. However, the investigation of neural responses to continuous dynamic input poses the challenge of how to link neural responses back to continuous sensory input. In the present tutorial, we provide a step-by-step introduction to one approach to tackle this issue, namely the use of linear models to investigate neural tracking responses in electroencephalographic (EEG) data. While neural tracking has gained increasing popularity in adult cognitive neuroscience over the past decade, its application to infant EEG is still rare and comes with its own challenges. After introducing the concept of neural tracking, we discuss and compare the use of forward vs. backward models and individual vs. generic models using an example data set of infant EEG data. Each section comprises a theoretical introduction as well as a concrete example using MATLAB code. We argue that neural tracking provides a promising way to investigate early (social) processing in an ecologically valid setting.

Re-directing attention to objects in working memory can enhance their representational fidelity. However, how this attentional enhancement of memory representations is implemented across distinct, sensory and cognitive-control brain network is unspecified. The present fMRI experiment leverages psychophysical modelling and multivariate auditory-pattern decoding as behavioral and neural proxies of mnemonic fidelity. Listeners performed an auditory syllable pitch-discrimination task and received retro-active cues to selectively attend to a to-be-probed syllable in memory. Accompanied by increased neural activation in fronto-parietal and cingulo-opercular networks, valid retro-cues yielded faster and more perceptually sensitive responses in recalling acoustic detail of memorized syllables. Information about the cued auditory object was decodable from hemodynamic response patterns in superior temporal sulcus (STS), fronto-parietal, and sensorimotor regions. However, among these regions retaining auditory memory objects, neural fidelity in the left STS and its enhancement through attention-to-memory best predicted individuals’ gain in auditory memory recall precision. Our results demonstrate how functionally discrete brain regions differentially contribute to the attentional enhancement of memory representations.

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.

Familiar and unfamiliar voice perception are often understood as being distinct from each other. For identity perception, theoretical work has proposed that listeners use acoustic information in different ways to perceive identity from familiar and unfamiliar voices: Unfamiliar voices are thought to be processed based on close comparisons of acoustic properties, while familiar voices are processed based on diagnostic acoustic features that activate a stored person-specific representation of that voice. To date no empirical study has directly examined whether and how familiar and unfamiliar listeners differ in their use of acoustic information for identity perception. Here, we tested this theoretical claim by linking listeners' judgements in voice identity tasks to complex acoustic representation — spectral similarity of the heard voice recordings. Participants (N = 177) who were either familiar or unfamiliar with a set of voices completed an identity discrimination task (Experiment 1) or an identity sorting task (Experiment 2). In both experiments, identity judgements for familiar and unfamiliar voices were guided by spectral similarity: Pairs of recordings with greater acoustic similarity were more likely to be perceived as belonging to the same voice identity. However, while there were no differences in how familiar and unfamiliar listeners used acoustic information for identity discrimination, differences were apparent for identity sorting. Our study therefore challenges proposals that view familiar and unfamiliar voice perception as being at all times distinct. Instead, our data suggest a critical role of the listening situation in which familiar and unfamiliar voices are evaluated, thus characterising voice identity perception as a highly dynamic process in which listeners opportunistically make use of any kind of information they can access.

Recent research posits that the cognitive system samples target stimuli in a rhythmic fashion, characterized by target detection fluctuating at frequencies of ~3–8 Hz. Besides prioritized encoding of targets, a key cognitive function is the protection of working memory from distractor intrusion. Here, we test to which degree the vulnerability of working memory to distraction is rhythmic. In an Irrelevant-Speech Task, N = 23 human participants had to retain the serial order of nine numbers in working memory while being distracted by task-irrelevant speech with variable temporal onsets. The magnitude of the distractor-evoked N1 component in the event-related potential as well as behavioural recall accuracy, both measures of memory distraction, were periodically modulated by distractor onset time in approximately 2–4 cycles per second (Hz). Critically, an underlying 2.5-Hz rhythm explained variation in both measures of distraction such that stronger phasic distractor encoding mediated lower phasic memory recall accuracy. In a behavioural follow-up experiment, we tested whether these results would replicate in a task design without rhythmic presentation of target items. Participants (N = 6 with on average >2,500 trials, each) retained two line-figures in memory while being distracted by acoustic noise of varying onset across trials. In agreement with the main experiment, the temporal onset of the distractor periodically modulated memory performance. These results suggest that during working memory retention, the human cognitive system implements distractor suppression in a temporally dynamic fashion, reflected in ~400-ms long cycles of high versus low distractibility.

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.

Anticipation of an impending stimulus shapes the state of the sensory systems, optimizing neural and behavioral responses. Here, we studied the role of brain oscillations in mediating spatial and temporal anticipations. Because spatial attention and temporal expectation are often associated with visual and auditory processing, respectively, we directly contrasted the visual and auditory modalities and asked whether these anticipatory mechanisms are similar in both domains. We recorded the magnetoencephalogram in healthy human participants performing an auditory and visual target discrimination task, in which cross-modal cues provided both temporal and spatial information with regard to upcoming stimulus presentation. Motivated by prior findings, we were specifically interested in delta (1–3 Hz) and alpha (8–13 Hz) band oscillatory state in anticipation of target presentation and their impact on task performance. Our findings support the view that spatial attention has a stronger effect in the visual domain, whereas temporal expectation effects are more prominent in the auditory domain. For the spatial attention manipulation, we found a typical pattern of alpha lateralization in the visual system, which correlated with response speed. Providing a rhythmic temporal cue led to increased postcue synchronization of low-frequency rhythms, although this effect was more broadband in nature, suggesting a general phase reset rather than frequency-specific neural entrainment. In addition, we observed delta-band synchronization with a frontal topography, which correlated with performance, especially in the auditory task. Combined, these findings suggest that spatial and temporal anticipations operate via a top–down modulation of the power and phase of low-frequency oscillations, respectively.

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 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.

Magneto- and electroencephalographic (M/EEG) signals recorded from the human scalp have allowed for substantial advances for neural models of speech comprehension over the past decades. These methods are currently advancing rapidly and continue to offer unparalleled insight in the near-to-real-time neural dynamics of speech processing. We provide a historically informed overview over dependent measures in the time and frequency domain and highlight recent advances resulting from these measures. We discuss the notorious challenges (and solutions) speech and language researchers are faced with when studying auditory brain responses in M/EEG. We argue that a key to understanding the neural basis of speech comprehension will lie in studying interactions between the neural tracking of speech and the functional neural network dynamics. This article is intended for both, non-experts who want to learn how to use M/EEG to study speech comprehension and scholars aiming for an overview of state-of-the-art M/...

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.

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.

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.

[This corrects the article DOI: 10.1371/journal.pbio.2002794.].

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.

Electroencephalography (EEG) continues to be the most popular method to investigate cognitive brain mechanisms in young children and infants. Most infant studies rely on the well-established and easy-to-use event-related brain potential (ERP). As a severe disadvantage, ERP computation requires a large number of repetitions of items from the same stimulus-category, compromising both ERPs’ reliability and their ecological validity in infant research. We here explore a way to investigate infant continuous EEG responses to an ongoing, engaging signal (i.e., “neural tracking”) by using multivariate temporal response functions (mTRFs), an approach increasingly popular in adult-EEG research. N=52 infants watched a 5-min episode of an age-appropriate cartoon while the EEG signal was recorded. We estimated and validated forward encoding models of auditory-envelope and visual-motion features. We compared individual and group-based (‘generic’) models of the infant brain response to comparison data from N=28 adults. The generic model yielded clearly defined response functions for both, the auditory and the motion regressor. Importantly, this response profile was present also on an individual level, albeit with lower precision of the estimate but above-chance predictive accuracy for the modelled individual brain responses. In sum, we demonstrate that mTRFs are a feasible way of analyzing continuous EEG responses in infants. We observe robust response estimates both across and within participants from only five minutes of recorded EEG signal. Our results open ways for incorporating more engaging and more ecologically valid stimulus materials when probing cognitive, perceptual, and affective processes in infants and young children.

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.

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.

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.

Children's language skills develop rapidly with increasing age, and several studies indicate that they use language- and age-specific strategies to understand complex sentences. In the present experiment, functional magnetic resonance imaging (fMRI) and behavioral measures were used to investigate the acquisition of case-marking cues for sentence interpretation in the developing brain of German preschool children with a mean age of 6 years. Short sentences were presented auditorily, consisting of a transitive verb and two case-marked arguments with canonical subject-initial or non canonical object-initial word order. Overall group results revealed mainly left hemispheric activation in the perisylvian cortex with increased activation in the inferior parietal cortex (IPC), and the anterior cingulate cortex (ACC) for object-initial compared to subject-initial sentences. However, single-subject analysis suggested two distinct activation patterns within the group which allowed a classification into two subgroups. One subgroup showed the predicted activation increase in the left inferior frontal gyrus (IFG) for the more difficult object-initial compared to subject-initial sentences, while the other group showed the reverse effect. This activation in the left IFG can be taken to reflect the degree to which adult-like sentence processing strategies, necessary to integrate case-marking information, are applied. Additional behavioral data on language development tests show that these two subgroups differ in their grammatical knowledge. Together with these behavioral findings, the results indicate that the use of a particular processing strategy is not dependent on age as such, but rather on the child's individual grammatical knowledge and the ability to use specific language cues for successful sentence comprehension.

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.

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.

Speech perception requires the decoding of complex acoustic patterns. According to most cognitive models of spoken word recognition, this complexity is dealt with before lexical access via a process of abstraction from the acoustic signal to pre-lexical categories. It is currently unclear how these categories are implemented in the auditory cortex. Recent advances in animal neurophysiology and human functional imaging have made it possible to investigate the processing of speech in terms of probabilistic cortical maps rather than simple cognitive subtraction, which will enable us to relate neurometric data more directly to behavioural studies. We suggest that integration of insights from cognitive science, neurophysiology and functional imaging is necessary for furthering our understanding of pre-lexical abstraction in the cortex.

Speech processing in auditory cortex and beyond is a remarkable yet poorly understood faculty of the listening brain. Here we show that stop consonants, as the most transient constituents of speech, are sufficient to involve speech perception circuits in the human superior temporal cortex. Left anterolateral superior temporal cortex showed a stronger response in blood oxygenation level--dependent functional magnetic resonance imaging (fMRI) to intelligible consonantal bursts compared with incomprehensible control sounds matched for spectrotemporal complexity. Simultaneously, the left posterior superior temporal plane (including planum temporale [PT]) exhibited a noncategorical responsivity to complex stimulus acoustics across all trials, showing no preference for intelligible speech sounds. Multistage hierarchical processing of speech sounds is thus revealed with fMRI, providing evidence for a role of the PT in the fundamental stages of the acoustic analysis of complex sounds, including speech.

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.

This editorial accompanies a special issue of Brain and Language re-visiting old themes and new leads in the electrophysiology of language. The event-related potential (ERP) as a series of characteristic deflections ("components") over time and their distribution on the scalp has been exploited by speech and language researchers over decades to find support for diverse psycholinguistic models. Fortunately, methodological and statistical advances have allowed human neuroscience to move beyond some of the limitations imposed when looking at the ERP only. Most importantly, we currently witness a refined and refreshed look at "event-related" (in the literal sense) brain activity that relates itself more closely to the actual neurobiology of speech and language processes. It is this imminent change in handling and interpreting electrophysiological data of speech and language experiments that this special issue intends to capture.

Meaningful auditory stimuli such as speech and music often vary simultaneously along multiple time scales. Thus, listeners must selectively attend to, and selectively ignore, separate but intertwined temporal features. The current study aimed to identify and characterize the neural network specifically involved in this feature-selective attention to time. We used a novel paradigm where listeners judged either the duration or modulation rate of auditory stimuli, and in which the stimulation, working memory demands, response requirements, and task difficulty were held constant. A first analysis identified all brain regions where individual brain activation patterns were correlated with individual behavioral performance patterns, which thus supported temporal judgments generically. A second analysis then isolated those brain regions that specifically regulated selective attention to temporal features: Neural responses in a bilateral fronto-parietal network including insular cortex and basal ganglia decreased with degree of change of the attended temporal feature. Critically, response patterns in these regions were inverted when the task required selectively ignoring this feature. The results demonstrate how the neural analysis of complex acoustic stimuli with multiple temporal features depends on a fronto-parietal network that simultaneously regulates the selective gain for attended and ignored temporal features.

The Arcuate Fasciculus/Superior Longitudinal Fasciculus (AF/SLF) is the white-matter bundle that connects posterior superior temporal and inferior frontal cortex. Its causal functional role in sentence processing and verbal working memory is currently under debate. While impairments of sentence processing and verbal working memory often co-occur in patients suffering from AF/SLF damage, it is unclear whether these impairments result from shared white-matter damage to the verbal-working-memory network. The present study sought to specify the behavioral consequences of focal AF/SLF damage for sentence processing and verbal working memory, which were assessed in a single patient suffering from a cleft-like lesion spanning the deep left superior temporal gyrus, sparing most surrounding gray matter. While tractography suggests that the ventral fronto-temporal white-matter bundle is intact in this patient, the AF/SLF was not visible to tractography. In line with the hypothesis that the AF/SLF is causally involved in sentence processing, the patient׳s performance was selectively impaired on sentences that jointly involve both complex word orders and long word-storage intervals. However, the patient was unimpaired on sentences that only involved long word-storage intervals without involving complex word orders. On the contrary, the patient performed generally worse than a control group across standard verbal-working-memory tests. We conclude that the AF/SLF not only plays a causal role in sentence processing, linking regions of the left dorsal inferior frontal gyrus to the temporo-parietal region, but moreover plays a crucial role in verbal working memory, linking regions of the left ventral inferior frontal gyrus to the left temporo-parietal region. Together, the specific sentence-processing impairment and the more general verbal-working-memory impairment may imply that the AF/SLF subserves both sentence processing and verbal working memory, possibly pointing to the AF and SLF respectively supporting each.

Auditory categorization is a vital skill involving the attribution of meaning to acoustic events, engaging domain-specific (i.e., auditory) as well as domain-general (e.g., executive) brain networks. A listener's ability to categorize novel acoustic stimuli should therefore depend on both, with the domain-general network being particularly relevant for adaptively changing listening strategies and directing attention to relevant acoustic cues. Here we assessed adaptive listening behavior, using complex acoustic stimuli with an initially salient (but later degraded) spectral cue and a secondary, duration cue that remained nondegraded. We employed voxel-based morphometry (VBM) to identify cortical and subcortical brain structures whose individual neuroanatomy predicted task performance and the ability to optimally switch to making use of temporal cues after spectral degradation. Behavioral listening strategies were assessed by logistic regression and revealed mainly strategy switches in the expected direction, with considerable individual differences. Gray-matter probability in the left inferior parietal lobule (BA 40) and left precentral gyrus was predictive of "optimal" strategy switch, while gray-matter probability in thalamic areas, comprising the medial geniculate body, co-varied with overall performance. Taken together, our findings suggest that successful auditory categorization relies on domain-specific neural circuits in the ascending auditory pathway, while adaptive listening behavior depends more on brain structure in parietal cortex, enabling the (re)direction of attention to salient stimulus properties. ? 2013 Published by Elsevier Ltd.

Our brain has the capacity of providing an experience of hearing even in the absence of auditory stimulation. This can be seen as illusory conscious perception. While increasing evidence postulates that conscious perception requires specific brain states that systematically relate to specific patterns of oscillatory activity, the relationship between auditory illusions and oscillatory activity remains mostly unexplained. To investigate this we recorded brain activity with magnetoencephalography and collected intracranial data from epilepsy patients while participants listened to familiar as well as unknown music that was partly replaced by sections of pink noise. We hypothesized that participants have a stronger experience of hearing music throughout noise when the noise sections are embedded in familiar compared to unfamiliar music. This was supported by the behavioral results showing that participants rated the perception of music during noise as stronger when noise was presented in a familiar context. Time-frequency data show that the illusory perception of music is associated with a decrease in auditory alpha power pointing to increased auditory cortex excitability. Furthermore, the right auditory cortex is concurrently synchronized with the medial temporal lobe, putatively mediating memory aspects associated with the music illusion. We thus assume that neuronal activity in the highly excitable auditory cortex is shaped through extensive communication between the auditory cortex and the medial temporal lobe, thereby generating the illusion of hearing music during noise.

Native listeners make use of higher-level, context-driven semantic and linguistic information during the perception of speech-in-noise. In a recent behavioral study, using a new paradigm that isolated the semantic level of speech by using words, we showed that this native-language benefit is at least partly driven by semantic context (Golestani et al., 2009). Here, we used the same paradigm in a functional magnetic resonance imaging (fMRI) experiment to study the neural bases of speech intelligibility, as well as to study the neural bases of this semantic context effect in the native language. A forced-choice recognition task on the first of two auditorily presented semantically related or unrelated words was employed, where the first, 'target' word was embedded in different noise levels. Results showed that activation in components of the brain language network, including Broca's area and the left posterior superior temporal sulcus, as well as brain regions known to be functionally related to attention and task difficulty, was modulated by stimulus intelligibility. In line with several previous studies examining the role of linguistic context in the intelligibility of degraded speech at the sentence level, we found that activation in the angular gyrus of the left inferior parietal cortex was modulated by the presence of semantic context, and further, that this modulation depended on the intelligibility of the speech stimuli. Our findings help to further elucidate neural mechanisms underlying the interaction of context-driven and signal-driven factors during the perception of degraded speech, and this specifically at the semantic level. (c) 2013 Elsevier Inc. All rights reserved.

In sentence processing, storage and ordering of the verb and its arguments (subject and object) are core tasks. Their cortical representation is a matter of ongoing debate, and it is unclear whether prefrontal activations in neuroimaging studies on sentence processing reflect the storage of arguments or their ordering. Moreover, it is unclear how storage during sentence processing relates to the neuroanatomy of storage outside the sentence processing domain. To tackle these questions, we crossed the factor “ordering” (subject-first vs. object-first German sentences) with the factor “storage” (one vs. four phrases intervene between the critical argument and the verb) in an auditory fMRI study. Ordering focally activated the left pars opercularis in Broca's area, while storage activated deep left temporo-parietal (TP) regions. Notably, left TP activation correlated with listener's digit span, while Broca's area activation did not. Furthermore, fractional anisotropy of listeners' left arcuate fasciculus/superior longitudinal fasciculus (AF/SLF) is shown to covary with the functional effect of increased storage demands at sites along the tract. Functionally, the results suggest that storage during sentence processing relies on TP regions, likely shared between sentence processing and other working memory-related tasks, while Broca's area appears as a distinct neural correlate of ordering. We conclude that the abstract notion of sentence processing can be captured by the interplay of concrete cognitive concepts such as ordering and storage.

Noise-vocoded speech is a spectrally highly degraded signal, but it preserves the temporal envelope of speech. Listeners vary considerably in their ability to adapt to this degraded speech signal. Here, we hypothesised that individual differences in adaptation to vocoded speech should be predictable by non-speech auditory, cognitive, and neuroanatomical factors. We tested 18 normal-hearing participants in a short-term vocoded speech-learning paradigm (listening to 100 4-band-vocoded sentences). Non-speech auditory skills were assessed using amplitude modulation (AM) rate discrimination, where modulation rates were centred on the speech-relevant rate of 4 Hz. Working memory capacities were evaluated (digit span and nonword repetition), and structural MRI scans were examined for anatomical predictors of vocoded speech learning using voxel-based morphometry. Listeners who learned faster to understand degraded speech also showed smaller thresholds in the AM discrimination task. This ability to adjust to degraded speech is furthermore reflected anatomically in increased grey matter volume in an area of the left thalamus (pulvinar) that is strongly connected to the auditory and prefrontal cortices. Thus, individual non-speech auditory skills and left thalamus grey matter volume can predict how quickly a listener adapts to degraded speech.

Modulations of human alpha oscillations (8–13 Hz) accompany many cognitive processes, but their functional role in auditory perception has proven elusive: Do oscillatory dynamics of alpha reflect acoustic details of the speech signal and are they indicative of comprehension success? Acoustically presented words were degraded in acoustic envelope and spectrum in an orthogonal design, and electroencephalogram responses in the frequency domain were analyzed in 24 participants, who rated word comprehensibility after each trial. First, the alpha power suppression during and after a degraded word depended monotonically on spectral and, to a lesser extent, envelope detail. The magnitude of this alpha suppression exhibited an additional and independent influence on later comprehension ratings. Second, source localization of alpha suppression yielded superior parietal, prefrontal, as well as anterior temporal brain areas. Third, multivariate classification of the time–frequency pattern across participants showed that patterns of late posterior alpha power allowed best for above-chance classification of word intelligibility. Results suggest that both magnitude and topography of late alpha suppression in response to single words can indicate a listener's sensitivity to acoustic features and the ability to comprehend speech under adverse listening conditions.

Under real-life adverse listening conditions, the interdependence of the brain's analysis of language structure (syntax) and its analysis of the acoustic signal is unclear. In two fMRI experiments, we first tested the functional neural organization when listening to increasingly complex syntax in fMRI. We then tested parametric combinations of syntactic complexity (argument scrambling in three degrees) with speech signal degradation (noise-band vocoding in three different numbers of bands), to shed light on the mutual dependency of sound and syntax analysis along the neural processing pathways. The left anterior and the posterior superior temporal sulcus (STS) as well as the left inferior frontal cortex (IFG) were linearly more activated as syntactic complexity increased (Experiment 1). In Experiment 2, when syntactic complexity was combined with improving signal quality, this pattern was replicated. However, when syntactic complexity was additive to degrading signal quality, the syntactic complexity effect in the IFG shifted dorsally and medially, and the activation effect in the left posterior STS shifted from posterior toward more middle sections of the sulcus. A distribution analysis of supra- as well as subthreshold data was indicative of this pattern of shifts in the anterior and posterior STS and within the IFG. Results suggest a signal quality gradient within the fronto-temporal language network. More signal-bound processing areas, lower in the processing hierarchy, become relatively more recruited for the analysis of complex language input under more challenging acoustic conditions (“upstream delegation”). This finding provides evidence for dynamic resource assignments along the neural pathways in auditory language comprehension.

Iconic gestures are spontaneous hand movements that illustrate certain contents of speech and, as such, are an important part of face-to-face communication. This experiment targets the brain bases of how iconic gestures and speech are integrated during comprehension. Areas of integration were identified on the basis of two classic properties of multimodal integration, bimodal enhancement and inverse effectiveness (i.e., greater enhancement for unimodally least effective stimuli). Participants underwent fMRI while being presented with videos of gesture-supported sentences as well as their unimodal components, which allowed us to identify areas showing bimodal enhancement. Additionally, we manipulated the signal-to-noise ratio of speech (either moderate or good) to probe for integration areas exhibiting the inverse effectiveness property. Bimodal enhancement was found at the posterior end of the superior temporal sulcus and adjacent superior temporal gyrus (pSTS/STG) in both hemispheres, indicating that the integration of iconic gestures and speech takes place in these areas. Furthermore, we found that the left pSTS/STG specifically showed a pattern of inverse effectiveness, i.e., the neural enhancement for bimodal stimulation was greater under adverse listening conditions. This indicates that activity in this area is boosted when an iconic gesture accompanies an utterance that is otherwise difficult to comprehend. The neural response paralleled the behavioral data observed. The present data extends results from previous gesture-speech integration studies in showing that pSTS/STG plays a key role in the facilitation of speech comprehension through simultaneous gestural input.