Individual prediction tendencies do not generalize across modalities.

Predictive processing theories, which model the brain as a "prediction machine", explain a wide range of cognitive functions, including learning, perception and action. Furthermore, it is increasingly accepted that aberrant prediction tendencies play a crucial role in psychiatric disorders. Given this explanatory value for clinical psychiatry, prediction tendencies are often implicitly conceptualized as individual traits or as tendencies that generalize across situations. As this has not yet explicitly been shown, in the current study, we quantify to what extent the individual tendency to anticipate sensory features of high probability generalizes across modalities. Using magnetoencephalography (MEG), we recorded brain activity while participants were presented with a sequence of four different (either visual or auditory) stimuli, which changed according to predefined transitional probabilities of two entropy levels: ordered vs. random. Our results show that, on a group-level, under conditions of low entropy, stimulus features of high probability are preactivated in the auditory but not in the visual modality. Crucially, the magnitude of the individual tendency to predict sensory events seems not to correlate between the two modalities. Furthermore, reliability statistics indicate poor internal consistency, suggesting that the measures from the different modalities are unlikely to reflect a single, common cognitive process. In sum, our findings suggest that quantification and interpretation of individual prediction tendencies cannot be generalized across modalities.

Neural speech tracking shifts from the syllabic to the modulation rate of speech as intelligibility decreases.

The most prominent acoustic features in speech are intensity modulations, represented by the amplitude envelope of speech. Synchronization of neural activity with these modulations supports speech comprehension. As the acoustic modulation of speech is related to the production of syllables, investigations of neural speech tracking commonly do not distinguish between lower-level acoustic (envelope modulation) and higher-level linguistic (syllable rate) information. Here we manipulated speech intelligibility using noise-vocoded speech and investigated the spectral dynamics of neural speech processing, across two studies at cortical and subcortical levels of the auditory hierarchy, using magnetoencephalography. Overall, cortical regions mostly track the syllable rate, whereas subcortical regions track the acoustic envelope. Furthermore, with less intelligible speech, tracking of the modulation rate becomes more dominant. Our study highlights the importance of distinguishing between envelope modulation and syllable rate and provides novel possibilities to better understand differences between auditory processing and speech/language processing disorders.

Involuntary shifts of spatial attention contribute to distraction-Evidence from oscillatory alpha power and reaction time data.

Imagine you are focusing on the traffic on a busy street to ride your bike safely when suddenly you hear the siren of an ambulance. This unexpected sound involuntarily captures your attention and interferes with ongoing performance. We tested whether this type of distraction involves a spatial shift of attention. We measured behavioral data and magnetoencephalographic alpha power during a cross-modal paradigm that combined an exogenous cueing task and a distraction task. In each trial, a task-irrelevant sound preceded a visual target (left or right). The sound was usually the same animal sound (i.e., standard sound). Rarely, it was replaced by an unexpected environmental sound (i.e., deviant sound). Fifty percent of the deviants occurred on the same side as the target, and 50% occurred on the opposite side. Participants responded to the location of the target. As expected, responses were slower to targets that followed a deviant compared to a standard. Crucially, this distraction effect was mitigated by the spatial relationship between the targets and the deviants: responses were faster when targets followed deviants on the same versus different side, indexing a spatial shift of attention. This was further corroborated by a posterior alpha power modulation that was higher in the hemisphere ipsilateral (vs. contralateral) to the location of the attention-capturing deviant. We suggest that this alpha power lateralization reflects a spatial attention bias. Overall, our data support the contention that spatial shifts of attention contribute to deviant distraction.

Gender differentiates effects of acoustic stimulation in patients with tinnitus.

Gender constitutes a major factor to consider when tailoring subtype-based therapies for tinnitus. Previous reports showed important differences between men and women concerning basic perceptual tinnitus characteristics (i.e., laterality, frequency, tinnitus loudness) as well as psychological reactions linked to this condition. Therapeutic approaches based on acoustic stimulation involve processes beyond a pure masking effect and consist of sound presentation temporarily altering or alleviating tinnitus perception via residual and/or lateral inhibition mechanisms. Presented stimuli may include pure tones, noise, and music adjusted to or modulated to filter out tinnitus pitch and therefore trigger reparative functional and structural changes in the auditory system. Furthermore, recent findings suggest that in tonal tinnitus, the presentation of pitch-adjusted sounds which were altered by a 10Hz modulation of amplitude was more efficient than unmodulated stimulation. In this paper, we investigate sex differences in the outcome of different variants of acoustic stimulation, looking for factors revealing predictive value in the efficiency of tinnitus relief.

Differential attention-dependent adjustment of frequency, power and phase in primary sensory and frontoparietal areas.

Continuously prioritizing behaviourally relevant information from the environment for improved stimulus processing is a crucial function of attention. In the current MEG study, we investigated how ongoing oscillatory activity of both sensory and non-sensory brain regions are differentially impacted by attentional focus. Low-frequency phase alignment of neural activity in primary sensory areas, with respect to attended/ignored features has been suggested to support top-down prioritization. However, phase adjustment in frontoparietal regions has not been widely studied, despite general implication of these in top-down selection of information. To investigate this, we let participants perform an established intermodal selective attention task, where low-frequency auditory (1.6 Hz) and visual (1.8 Hz) stimuli were presented simultaneously. We instructed them to either attend to the auditory or to the visual stimuli and to detect targets while ignoring the other stimulus stream. As expected, the strongest phase adjustment was observed in primary sensory regions for auditory and for visual stimulation, independent of attentional focus. We found greater differences in phase locking between attended and ignored stimulation for the visual modality. Interestingly, auditory temporal regions show small but significant attention-dependent neural entrainment even for visual stimulation. Extending findings from invasive recordings in non-human primates, we demonstrate an effect of attentional focus on the phase of the entrained oscillations in auditory and visual cortex which may be driven by phase locked increases of induced power. While sensory areas adjusted the phase of the respective stimulation frequencies, attentional focus adjusted the peak frequencies in nonsensory areas. Spatially these areas show a striking overlap with core regions of the dorsal attention network and the frontoparietal network. This suggests that these areas prioritize the attended modality by optimally exploiting the temporal structure of stimulation. Overall, our study complements and extends previous work by showing a differential effect of attentional focus on entrained oscillations (or phase adjustment) in primary sensory areas and frontoparietal areas.

Auditory cortical alpha/beta desynchronization prioritizes the representation of memory items during a retention period.

To-be-memorized information in working-memory could be protected against distracting influences by processes of functional inhibition or prioritization. Modulations of oscillations in the alpha to beta range in task-relevant sensory regions have been suggested to play an important role for both mechanisms. We adapted a Sternberg task variant to the auditory modality, with a strong or a weak distracting sound presented at a predictable time during the retention period. Using a time-generalized decoding approach, relatively decreased strength of memorized information was found prior to strong distractors, paralleled by decreased pre-distractor alpha/beta power in the left superior temporal gyrus (lSTG). Over the entire group, reduced beta power in lSTG was associated with relatively increased strength of memorized information. The extent of alpha power modulations within participants was negatively correlated with strength of memorized information. Overall, our results are compatible with a prioritization account, but point to nuanced differences between alpha and beta oscillations.

A backward encoding approach to recover subcortical auditory activity.

Several subcortical nuclei along the auditory pathway are involved in the processing of sounds. One of the most commonly used methods of measuring the activity of these nuclei is the auditory brainstem response (ABR). Due to its low signal-to-noise ratio, ABR's have to be derived by averaging over activity generated by thousands of artificial sounds such as clicks or tone bursts. This approach cannot be easily applied to natural listening situations (e.g. speech, music), which limits auditory cognitive neuroscientific studies to investigate mostly cortical processes. We propose that by individually training backward encoding models to reconstruct evoked ABRs from high-density electrophysiological data, spatial filters can be tuned to auditory brainstem activity. Since these individualized filters can be applied (i.e. generalized) to any other data set using the same spatial coverage, this could allow for the estimation of auditory brainstem activity from any continuous sensor level data. In this study, we established a proof-of-concept by using backward encoding models generated using a click stimulation rate of 30 â€‹Hz to predict ABR activity recorded using EEG from an independent measurement using a stimulation rate of 9 â€‹Hz. We show that individually predicted and measured ABR's are highly correlated (r â€‹~ â€‹0.7). Importantly these predictions are stable even when applying the trained backward encoding model to a low number of trials, mimicking a situation with an unfavorable signal-to-noise ratio. Overall, this work lays the necessary foundation to use this approach in more interesting listening situations.

Decoding across sensory modalities reveals common supramodal signatures of conscious perception.

An increasing number of studies highlight common brain regions and processes in mediating conscious sensory experience. While most studies have been performed in the visual modality, it is implicitly assumed that similar processes are involved in other sensory modalities. However, the existence of supramodal neural processes related to conscious perception has not been convincingly shown so far. Here, we aim to directly address this issue by investigating whether neural correlates of conscious perception in one modality can predict conscious perception in a different modality. In two separate experiments, we presented participants with successive blocks of near-threshold tasks involving subjective reports of tactile, visual, or auditory stimuli during the same magnetoencephalography (MEG) acquisition. Using decoding analysis in the poststimulus period between sensory modalities, our first experiment uncovered supramodal spatiotemporal neural activity patterns predicting conscious perception of the feeble stimulation. Strikingly, these supramodal patterns included activity in primary sensory regions not directly relevant to the task (e.g., neural activity in visual cortex predicting conscious perception of auditory near-threshold stimulation). We carefully replicate our results in a control experiment that furthermore show that the relevant patterns are independent of the type of report (i.e., whether conscious perception was reported by pressing or withholding a button press). Using standard paradigms for probing neural correlates of conscious perception, our findings reveal a common signature of conscious access across sensory modalities and illustrate the temporally late and widespread broadcasting of neural representations, even into task-unrelated primary sensory processing regions.

Stereotactic electroencephalography in humans reveals multisensory signal in early visual and auditory cortices.

Unequivocally demonstrating the presence of multisensory signals at the earliest stages of cortical processing remains challenging in humans. In our study, we relied on the unique spatio-temporal resolution provided by intracranial stereotactic electroencephalographic (SEEG) recordings in patients with drug-resistant epilepsy to characterize the signal extracted from early visual (calcarine and pericalcarine) and auditory (Heschl's gyrus and planum temporale) regions during a simple audio-visual oddball task. We provide evidences that both cross-modal responses (visual responses in auditory cortex or the reverse) and multisensory processing (alteration of the unimodal responses during bimodal stimulation) can be observed in intracranial event-related potentials (iERPs) and in power modulations of oscillatory activity at different temporal scales within the first 150 msec after stimulus onset. The temporal profiles of the iERPs are compatible with the hypothesis that MSI occurs by means of direct pathways linking early visual and auditory regions. Our data indicate, moreover, that MSI mainly relies on modulations of the low-frequency bands (foremost the theta band in the auditory cortex and the alpha band in the visual cortex), suggesting the involvement of feedback pathways between the two sensory regions. Remarkably, we also observed high-gamma power modulations by sounds in the early visual cortex, thus suggesting the presence of neuronal populations involved in auditory processing in the calcarine and pericalcarine region in humans.

Auditory cortical generators of the Frequency Following Response are modulated by intermodal attention.

The efferent auditory system suggests that brainstem auditory regions could also be sensitive to top-down processes. In electrophysiology, the Frequency Following Response (FFR) to speech stimuli has been used extensively to study brainstem areas. Despite seemingly straight-forward in addressing the issue of attentional modulations of brainstem regions by means of the FFR, the existing results are inconsistent. Moreover, the notion that the FFR exclusively represents subcortical generators has been challenged. We aimed to gain a more differentiated perspective on how the generators of the FFR are modulated by either attending to the visual or auditory input while neural activity was recorded using magnetoencephalography (MEG). In a first step our results confirm the strong contribution of also cortical regions to the FFR. Interestingly, of all regions exhibiting a measurable FFR response, only the right primary auditory cortex was significantly affected by intermodal attention. By showing a clear cortical contribution to the attentional FFR effect, our work significantly extends previous reports that focus on surface level recordings only. It underlines the importance of making a greater effort to disentangle the different contributing sources of the FFR and serves as a clear precaution of simplistically interpreting the FFR as brainstem response.

tACS-mediated modulation of the auditory steady-state response as seen with MEG.

BACKGROUND: Previous studies have shown that transcranial electrical stimulation can be successfully applied during simultaneous MEG measurements. In particular, using beamforming they have established that changes of stimulus induced as well as evoked activity can be inspected during transcranial alternating current stimulation (tACS). OBJECTIVE/HYPOTHESIS: We studied tACS-mediated changes of the auditory steady-state response (ASSR), hypothesizing that-due to the putatively inhibitory role of alpha oscillations-these evoked responses would be diminished. METHODS: We compared ASSRs in conditions with and without 12-Hz and 6.5-Hz sinusoidal 1.5 mA tACS, applied bilaterally over temporal areas. Source-level activity was estimated using a linearly constrained minimum variance beamformer and compared across tACS conditions using paired t-tests following a condition-internal normalization procedure. CONCLUSIONS: By separating the electrical and auditory stimulation to non-overlapping parts of the frequency spectrum, we were able to compare auditory-evoked steady-state activity across tACS conditions. We observed a significant decrease in normalized ASSR power in the 12-Hz tACS condition, illustrating that tACS could induce immediate changes in auditory evoked activity. This study sets a methodology to further interrogate the causal roles of oscillatory dynamics in auditory cortices, as well as suggests perspectives for employing tACS in clinical contexts.

A Visual Cortical Network for Deriving Phonological Information from Intelligible Lip Movements.

Successful lip-reading requires a mapping from visual to phonological information [1]. Recently, visual and motor cortices have been implicated in tracking lip movements (e.g., [2]). It remains unclear, however, whether visuo-phonological mapping occurs already at the level of the visual cortex-that is, whether this structure tracks the acoustic signal in a functionally relevant manner. To elucidate this, we investigated how the cortex tracks (i.e., entrains to) absent acoustic speech signals carried by silent lip movements. Crucially, we contrasted the entrainment to unheard forward (intelligible) and backward (unintelligible) acoustic speech. We observed that the visual cortex exhibited stronger entrainment to the unheard forward acoustic speech envelope compared to the unheard backward acoustic speech envelope. Supporting the notion of a visuo-phonological mapping process, this forward-backward difference of occipital entrainment was not present for actually observed lip movements. Importantly, the respective occipital region received more top-down input, especially from left premotor, primary motor, and somatosensory regions and, to a lesser extent, also from posterior temporal cortex. Strikingly, across participants, the extent of top-down modulation of the visual cortex stemming from these regions partially correlated with the strength of entrainment to absent acoustic forward speech envelope, but not to present forward lip movements. Our findings demonstrate that a distributed cortical network, including key dorsal stream auditory regions [3-5], influences how the visual cortex shows sensitivity to the intelligibility of speech while tracking silent lip movements.

Amplitude modulation rate dependent topographic organization of the auditory steady-state response in human auditory cortex.

Periodic modulations of an acoustic feature, such as amplitude over a certain frequency range, leads to phase locking of neural responses to the envelope of the modulation. Using electrophysiological methods this neural activity pattern, also called the auditory steady-state response (aSSR), is visible following frequency transformation of the evoked response as a clear spectral peak at the modulation frequency. Despite several studies employing the aSSR that show, for example, strongest responses for ∼40 Hz and an overall right-hemispheric dominance, it has not been investigated so far to what extent within auditory cortex different modulation frequencies elicit aSSRs at a homogenous source or whether the localization of the aSSR is topographically organized in a systematic manner. The latter would be suggested by previous neuroimaging works in monkeys and humans showing a periodotopic organization within and across distinct auditory fields. However, the sluggishness of the signal from these neuroimaging works prohibit inferences with regards to the fine-temporal features of the neural response. In the present study, we employed amplitude-modulated (AM) sounds over a range between 4 and 85 Hz to elicit aSSRs while recording brain activity via magnetoencephalography (MEG). Using beamforming and a fine spatially resolved grid restricted to auditory cortical processing regions, our study revealed a topographic representation of the aSSR that depends on AM rate, in particular in the medial-lateral (bilateral) and posterior-anterior (right auditory cortex) direction. In summary, our findings confirm previous studies that showing different AM rates to elicit maximal response in distinct neural populations. They extend these findings however by also showing that these respective neural ensembles in auditory cortex actually phase lock their activity over a wide modulation frequency range.

Cross-modal distractors modulate oscillatory alpha power: the neural basis of impaired task performance.

Unexpected novel sounds capture one's attention, even when irrelevant to the task pursued (e.g., playing video game). This often comes at a cost to the task (e.g., slower responding). The neural basis for this behavioral distraction effect is not well understood and is subject of this study. Our approach was motivated by findings from cuing paradigms suggesting a link between modulations in oscillatory activity and voluntary attention shifts. The current study tested whether oscillatory activity is also modulated by a task-irrelevant auditory distractor, reflecting a neural signature of an involuntary shift of attention and accounting for the impaired task performance. We reanalyzed magnetoencephalographic data collected via an auditory-visual distraction paradigm in which a task-relevant visual stimulus was preceded by a task-irrelevant sound on each trial. In 87.5% this was a regular sound (Standard); in 12.5% this was a novel sound (Distractor). We compared nonphase locked oscillatory activity in a time window prior to the visual target as a function of the experimental manipulation (Distractor, Standard). We found low power in the pretarget time window for Distractors compared to Standards in the alpha and beta frequency bands. Importantly, individual alpha power correlated with response speed on a trial-by-trial basis for the Distractor only. Sources were localized to the occipital cortex, and also to the parietal and supratemporal cortices. These findings support our hypothesis that the distractor-related alpha power modulation indexes an involuntary shift of attention which accounts for the impaired task performance.

Prestimulus Network Integration of Auditory Cortex Predisposes Near-Threshold Perception Independently of Local Excitability.

An ever-increasing number of studies are pointing to the importance of network properties of the brain for understanding behavior such as conscious perception. However, with regards to the influence of prestimulus brain states on perception, this network perspective has rarely been taken. Our recent framework predicts that brain regions crucial for a conscious percept are coupled prior to stimulus arrival, forming pre-established pathways of information flow and influencing perceptual awareness. Using magnetoencephalography (MEG) and graph theoretical measures, we investigated auditory conscious perception in a near-threshold (NT) task and found strong support for this framework. Relevant auditory regions showed an increased prestimulus interhemispheric connectivity. The left auditory cortex was characterized by a hub-like behavior and an enhanced integration into the brain functional network prior to perceptual awareness. Right auditory regions were decoupled from non-auditory regions, presumably forming an integrated information processing unit with the left auditory cortex. In addition, we show for the first time for the auditory modality that local excitability, measured by decreased alpha power in the auditory cortex, increases prior to conscious percepts. Importantly, we were able to show that connectivity states seem to be largely independent from local excitability states in the context of a NT paradigm.

Thalamocortical interactions underlying visual fear conditioning in humans.

Despite a strong focus on the role of the amygdala in fear conditioning, recent works point to a more distributed network supporting fear conditioning. We aimed to elucidate interactions between subcortical and cortical regions in fear conditioning in humans. To do this, we used two fearful faces as conditioned stimuli (CS) and an electrical stimulation at the left hand, paired with one of the CS, as unconditioned stimulus (US). The luminance of the CS was rhythmically modulated leading to "entrainment" of brain oscillations at a predefined modulation frequency. Steady-state responses (SSR) were recorded by MEG. In addition to occipital regions, spectral analysis of SSR revealed increased power during fear conditioning particularly for thalamus and cerebellum contralateral to the upcoming US. Using thalamus and amygdala as seed-regions, directed functional connectivity was calculated to capture the modulation of interactions that underlie fear conditioning. Importantly, this analysis showed that the thalamus drives the fusiform area during fear conditioning, while amygdala captures the more general effect of fearful faces perception. This study confirms ideas from the animal literature, and demonstrates for the first time the central role of the thalamus in fear conditioning in humans.

What it means to be Zen: marked modulations of local and interareal synchronization during open monitoring meditation.

Experienced meditators are able to voluntarily modulate their state of consciousness and attention. In the present study, we took advantage of this ability and studied brain activity related to the shift of mental state. Electrophysiological activity, i.e. EEG, was recorded from 11 subjects with varying degrees of meditation experience during Zen meditation (a form of open monitoring meditation) and during non-meditation rest. On a behavioral level, mindfulness scores were assessed using the Mindfulness Attention and Awareness Scale (MAAS). Analysis of EEG source power revealed the so far unreported finding that MAAS scores significantly correlated with gamma power (30-250Hz), particularly high-frequency gamma (100-245Hz), during meditation. High levels of mindfulness were related to increased high-frequency gamma, for example, in the cingulate cortex and somatosensory cortices. Further, we analyzed the relationship between connectivity during meditation and self-reported mindfulness (MAAS). We found a correlation between graph measures in the 160-170Hz range and MAAS scores. Higher levels of mindfulness were related to lower small worldedness as well as global and local clustering in paracentral, insular, and thalamic regions during meditation. In sum, the present study shows significant relationships of mindfulness and brain activity during meditation indicated by measures of oscillatory power and graph theoretical measures. The most prominent effects occur in brain structures crucially involved in processes of awareness and attention, which also show structural changes in short- and long-term meditators, suggesting continuative alterations in the meditating brain. Overall, our study reveals strong changes in ongoing oscillatory activity as well as connectivity patterns that appear to be sensitive to the psychological state changes induced by Zen meditation.

Listen to Yourself: The Medial Prefrontal Cortex Modulates Auditory Alpha Power During Speech Preparation.

How do we process stimuli that stem from the external world and stimuli that are self-generated? In the case of voice perception it has been shown that evoked activity elicited by self-generated sounds is suppressed compared with the same sounds played-back externally. We here wanted to reveal whether neural excitability of the auditory cortex-putatively reflected in local alpha band power--is modulated already prior to speech onset, and which brain regions may mediate such a top-down preparatory response. In the left auditory cortex we show that the typical alpha suppression found when participants prepare to listen disappears when participants expect a self-spoken sound. This suggests an inhibitory adjustment of auditory cortical activity already before sound onset. As a second main finding we demonstrate that the medial prefrontal cortex, a region known for self-referential processes, mediates these condition-specific alpha power modulations. This provides crucial insights into how higher-order regions prepare the auditory cortex for the processing of self-generated sounds. Furthermore, the mechanism outlined could provide further explanations to self-referential phenomena, such as "tickling yourself". Finally, it has implications for the so-far unsolved question of how auditory alpha power is mediated by higher-order regions in a more general sense.

Selective modulation of auditory cortical alpha activity in an audiovisual spatial attention task.

Despite substantial research on attentional modulations of visual alpha activity, doubts remain as to the existence and functional relevance of auditory cortical alpha-band oscillations. It has been argued that auditory cortical alpha does not exist, cannot be measured noninvasively, or that it is dependent on visual alpha generators. This study aimed to address these remaining doubts concerning auditory cortical alpha. A magnetoencephalography study was conducted using a combined audiovisual spatial cueing paradigm. In each trial, a cue indicated the side (left or right) and the modality (auditory or visual) to attend, followed by a short lateralized auditory or visual stimulus. Participants were instructed to respond to the stimuli by a button press. Results show that auditory cortical alpha power is selectively modulated by the audiospatial, but not the visuospatial, attention task. These findings provide further evidence for a distinct auditory cortical alpha generator, which can be measured noninvasively.

Synchronisation signatures in the listening brain: a perspective from non-invasive neuroelectrophysiology.

Human magneto- and electroencephalography (M/EEG) are capable of tracking brain activity at millisecond temporal resolution in an entirely non-invasive manner, a feature that offers unique opportunities to uncover the spatiotemporal dynamics of the hearing brain. In general, precise synchronisation of neural activity within as well as across distributed regions is likely to subserve any cognitive process, with auditory cognition being no exception. Brain oscillations, in a range of frequencies, are a putative hallmark of this synchronisation process. Embedded in a larger effort to relate human cognition to brain oscillations, a field of research is emerging on how synchronisation within, as well as between, brain regions may shape auditory cognition. Combined with much improved source localisation and connectivity techniques, it has become possible to study directly the neural activity of auditory cortex with unprecedented spatio-temporal fidelity and to uncover frequency-specific long-range connectivities across the human cerebral cortex. In the present review, we will summarise recent contributions mainly of our laboratories to this emerging domain. We present (1) a more general introduction on how to study local as well as interareal synchronisation in human M/EEG; (2) how these networks may subserve and influence illusory auditory perception (clinical and non-clinical) and (3) auditory selective attention; and (4) how oscillatory networks further reflect and impact on speech comprehension. This article is part of a Special Issue entitled Human Auditory Neuroimaging.