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It has been demonstrated that, while otherwise detrimental, noise can improve sensory perception under optimal conditions. The mechanism underlying this improvement is stochastic resonance. An inverted U-shaped relationship between noise level and task performance is considered as the signature of stochastic resonance. Previous studies have proposed the existence of stochastic resonance also in the human auditory system. However, the reported beneficial effects of noise are small, based on a small sample, and do not confirm the proposed inverted U-shaped function. Here, we investigated in two separate studies whether stochastic resonance may be present in the human auditory system by applying noise of different levels, either acoustically or electrically via transcranial random noise stimulation, while participants had to detect acoustic stimuli adjusted to their individual hearing threshold. We find no evidence for behaviorally relevant effects of stochastic resonance. Although detection rate for near-threshold acoustic stimuli appears to vary in an inverted U-shaped manner for some subjects, it varies in a U-shaped manner or in other manners for other subjects. Our results show that subjects do not benefit from noise, irrespective of its modality. In conclusion, our results question the existence of stochastic resonance in the human auditory system.

Interventions in developmental dyslexia typically consist of orthography-based reading and writing trainings. However, their efficacy is limited and, consequently, the symptoms persist into adulthood. Critical for this lack of efficacy is the still ongoing debate about the core deficit in dyslexia and its underlying neurobiological causes. There is ample evidence on phonological as well as auditory temporal processing deficits in dyslexia and, on the other hand, cortical gamma oscillations in the auditory cortex as functionally relevant for the extraction of linguistically meaningful information units from the acoustic signal. The present work aims to shed more light on the link between auditory gamma oscillations, phonological awareness, and literacy skills in dyslexia. By mean of EEG, individual gamma frequencies were assessed in a group of children and adolescents diagnosed with dyslexia as well as in an age-matched control group with typical literacy skills. Furthermore, phonological awareness was assessed in both groups, while in dyslexic participants also reading and writing performance was measured. We found significantly lower gamma peak frequencies as well as lower phonological awareness scores in dyslexic participants compared to age-matched controls. Additionally, results showed a positive correlation between the individual gamma frequency and phonological awareness. Our data suggest a hierarchical structure of neural gamma oscillations, phonological awareness, and literacy skills. Thereby, the results emphasize altered gamma oscillation not only as a core deficit in dyslexia but also as a potential target for future causal interventions. We discuss these findings considering non-invasive brain stimulation techniques and suggest transcranial alternating current stimulation as a promising approach to normalize dysfunctional oscillations in dyslexia.

Interventions in developmental dyslexia typically consist of orthography-based reading and writing trainings. However, their efficacy is limited and, consequently, the symptoms persist into adulthood. Critical for this lack of efficacy is the still ongoing debate about the core deficit in dyslexia and its underlying neurobiological causes. There is ample evidence on phonological as well as auditory temporal processing deficits in dyslexia and, on the other hand, cortical gamma oscillations in the auditory cortex as functionally relevant for the extraction of linguistically meaningful information units from the acoustic signal. The present work aims to shed more light on the link between auditory gamma oscillations, phonological awareness, and literacy skills in dyslexia. By mean of EEG, individual gamma frequencies were assessed in a group of children and adolescents diagnosed with dyslexia as well as in an age-matched control group with typical literacy skills. Furthermore, phonological awareness was assessed in both groups, while in dyslexic participants also reading and writing performance was measured. We found significantly lower gamma peak frequencies as well as lower phonological awareness scores in dyslexic participants compared to age-matched controls. Additionally, results showed a positive correlation between the individual gamma frequency and phonological awareness. Our data suggest a hierarchical structure of neural gamma oscillations, phonological awareness, and literacy skills. Thereby, the results emphasize altered gamma oscillation not only as a core deficit in dyslexia but also as a potential target for future causal interventions. We discuss these findings considering non-invasive brain stimulation techniques and suggest transcranial alternating current stimulation as a promising approach to normalize dysfunctional oscillations in dyslexia.

Most acoustic events in our environment do not appear randomly but are rather predictable due to the temporal regularity in that they occur. Besides sensory-related cortical areas, the cerebellum has been suggested as a key structure in temporal processing and in the anticipation of future events. Hence, patients with cerebellum lesions show impaired precision in temporal processing as reflected in the reduced ability to exploit temporal regularity. Using transcranial direct current stimulation (tDCS), we here aimed to draw further causal conclusions on the human cerebellum as functionally relevant in temporal processing of acoustic events. We focused on the electrophysiologic P3b, a large positive wave apparent in the electroencephalography (EEG), that represents encoding of task-relevant events and that has been demonstrated as sensitive to the exploitation of temporal regularities. Participants received 30 min of anodal, cathodal or sham tDCS over the cerebellum while they performed two oddball paradigms with different temporal regularities in that the acoustic stimuli were presented. Following clinical observations, we hypothesized that tDCS-effects will be present in the regular oddball paradigm only, thus, in the condition that allows anticipating the occurrence of subsequent stimuli. In result, we found that cathodal tDCS over the cerebellum reduced the P3b-amplitude specifically in response to target stimuli in the regular paradigm. Thereby, tDCS-induced changes mirror the effects of cerebellar lesions in clinical samples. Our data provides direct evidence for a causal link between the human cerebellum and auditory processing of temporal regularity and emphasize future work on a potential benefit of cerebellar-tDCS in clinical samples.

Selective attention is a basic process required to maintain goal-directed behavior by appropriately responding to target stimuli and suppressing reactions to non-target stimuli. It has been proposed that auditory selective attention is linked to the activity of the locus coeruleus-norepinergic (LC-NE) system and a large-scale fronto-parietal cortical network, but there is still sparse causal evidence for these assumptions. By applying transcutaneous vagal nerve stimulation (tVNS) and transcranial random noise stimulation (tRNS) over the frontal cortex, we systematically assessed the involvement of these subcortical and cortical components in the regulation of auditory selective attention. Using a single-blinded, sham-controlled, within-subject design we analyzed online effects of tVNS and tRNS in 20 healthy participants during an auditory oddball paradigm. We show significant stimulation-dependent modulations of auditory selective attention on the behavioral and electrophysiological level. Compared to sham, tVNS increased the P3 amplitude, while tRNS reduced the reaction time to target stimuli. Moreover, both techniques reduced the P3 latency. Our data provide evidence for the functional relevance of the subcortical NE system in the regulation of neural resources that allows a phasic response to incoming target stimuli. They indicate that frontal cortex structures are crucially involved in the successful evaluation of the respective information. Moreover, our results are in favor of the LC-P3 hypothesis claiming the vital role of the NE system in auditory selective attention and in the generation of the P3. Of note, the effects of tVNS on auditory selective attention are comparable with those evoked by pharmacological interventions and invasive vagal nerve stimulation.

Fatigue is one of the most common and debilitating symptoms affecting patients with multiple sclerosis (MS). Sustained cognitive effort induces cognitive fatigue, operationalized as subjective exhaustion and fatigue-related objective alertness decrements with time-on-task. During prolonged cognitive testing, MS patients show increased simple reaction times (RT) accompanied by lower amplitudes and prolonged latencies of the P300 event-related potential. Previous studies suggested a major role of structural and functional abnormalities in the frontal cortex including a frontal hypo-activation in fatigue pathogenesis. In the present study we investigated the neuromodulatory effect of transcranial direct current stimulation (tDCS) over the left dorsolateral prefrontal cortex (DLPFC) on objective measures of fatigue-related decrements in cognitive performance in MS patients. P300 during an auditory oddball task and simple reaction times in an alertness test were recorded at baseline, during and after stimulation. Compared to sham, anodal tDCS caused an increase in P300 amplitude that persisted after the end of stimulation and eliminated the fatigue-related increase in RT over the course of a testing session. Our findings demonstrate that anodal tDCS over the left DLPFC can counteract performance decrements associated with fatigue thereby leading to an improvement in the patient's ability to cope with sustained cognitive demands. This provides causal evidence for the functional relevance of the left DLPFC in fatigue pathophysiology. The results indicate that tDCS-induced modulations of frontal activity can be an effective therapeutic option for the treatment of fatigue-related declines in cognitive performance in MS patients.

Background One key mechanism thought to underlie speech processing is the alignment of cortical brain rhythms to the acoustic input, a mechanism termed entrainment. Recent work showed that transcranial electrical stimulation (tES) in speech relevant frequencies or adapted to the speech envelope can in fact enhance speech processing. However, it is unclear whether an oscillatory tES is necessary, or if transients in the stimulation (e.g., peaks in the tES signal) at relevant times are sufficient. Objective In this study we used a novel pulsed-tES-protocol and tested behaviorally if a transiently pulsed - instead of a persistently oscillating - tES signal, can improve speech processing. Methods While subjects listened to spoken sentences embedded in noise, brief electric direct current pulses aligned to speech transients (syllable onsets) were applied to auditory cortex regions to modulate comprehension. Additionally, we modulated the temporal delay between tES-pulses and speech transients to test for periodic modulations of behavior, indicative of entrainment by tES. Results Speech comprehension was improved when tES-pulses were applied with a delay of 100 ms in respect to the speech transients. Contradictory to previous reports we find no periodic modulation of behavior. However, we find indications that periodic modulations can be spurious results of sampling behavioral data too coarsely. Conclusions Subject’s speech comprehension benefits from pulsed-tES, yet behavior is not modulated periodically. Thus, pulsed-tES can aid cortical entrainment to speech input, which is especially relevant in a noisy environment. Yet, pulsed-tES does not seem to entrain brain oscillations by itself.

Background About 10% of the western population suffers from a specific disability in the acquisition of reading and writing skills, known as developmental dyslexia (DD). Even though DD starts in childhood it frequently continuous throughout lifetime. Impaired processing of acoustic features at the phonematic scale based on dysfunctional auditory temporal resolution is considered as one core deficit underlying DD. Recently, the efficacy of transcranial electrical stimulation (tES) to modulate auditory temporal resolution and phoneme processing in healthy individuals has been demonstrated. Objective The present work aims to investigate online effects of tES on phoneme processing in individuals with DD. Method Using an established phoneme-categorization task, we assessed the immediate behavioral and electrophysiological effects of transcranial alternating current stimulation (tACS) and transcranial random noise stimulation (tRNS) over bilateral auditory cortex in children and adolescents with DD (study 1) and adults with DD (study 2) on auditory phoneme processing acuity. Results Our data revealed that tACS improved phoneme categorization in children and adolescents with DD, an effect that was paralleled by an increase in evoked brain response patterns representing low-level sensory processing. In the adult sample we replicated these findings and additionally showed a more pronounced impact of tRNS on phoneme-categorization acuity. Conclusion These results provide compelling evidence for the potential of both tACS and tRNS to increase temporal precision of the auditory system in DD and suggest transcranial electrical stimulation as potential intervention in DD to foster the effect of standard phonology-based training.

The present study investigated the functional relevance of gamma oscillations for the processing of rapidly changing acoustic features in speech signals. For this purpose we analyzed repetition-induced perceptual learning effects in 18 healthy adult participants. The participants received either 6Hz or 40Hz tACS over the bilateral auditory cortex, while repeatedly performing a phoneme categorization task. In result, we found that 40Hz tACS led to a specific alteration in repetition-induced perceptual learning. While participants in the non-stimulated control group as well as those in the experimental group receiving 6Hz tACS considerably improved their perceptual performance, the application of 40Hz tACS selectively attenuated the repetition-induced improvement in phoneme categorization abilities. Our data provide causal evidence for a functional relevance of gamma oscillations during the perceptual learning of acoustic speech features. Moreover, we demonstrate that even less than twenty minutes of alternating current stimulation below the individual perceptual threshold is sufficient to affect speech perception. This finding is relevant in that this novel approach might have implications with respect to impaired speech processing in dyslexics and older adults.

Neural oscillations in the gamma range are the dominant rhythmic activation pattern in the human auditory cortex. These gamma oscillations are functionally relevant for the processing of rapidly changing acoustic information in both speech and non-speech sounds. Accordingly, there is a tight link between the temporal resolution ability of the auditory system and inherent neural gamma oscillations. Transcranial random noise stimulation (tRNS) has been demonstrated to specifically increase gamma oscillation in the human auditory cortex. However, neither the physiological mechanisms of tRNS nor the behavioral consequences of this intervention are completely understood. In the present study we stimulated the human auditory cortex bilaterally with tRNS while EEG was continuously measured. Modulations in the participants’ temporal and spectral resolution ability were investigated by means of a gap detection task and a pitch discrimination task. Compared to sham, auditory tRNS increased the detection rate for near-threshold stimuli in the temporal domain only, while no such effect was present for the discrimination of spectral features. Behavioral findings were paralleled by reduced peak latencies of the P50 and N1 component of the auditory event-related potentials (ERP) indicating an impact on early sensory processing. The facilitating effect of tRNS was limited to the processing of near-threshold stimuli while stimuli clearly below and above the individual perception threshold were not affected by tRNS. This non-linear relationship between the signal-to-noise level of the presented stimuli and the effect of stimulation further qualifies stochastic resonance (SR) as the underlying mechanism of tRNS on auditory processing. Our results demonstrate a tRNS related improvement in acoustic perception of time critical auditory information and, thus, provide further indices that auditory tRNS can amplify the resonance frequency of the auditory system.

Theoretical background Developmental dyslexia (DD) refers to the pathological impairment in the acquisition of reading and/or writing skills that is not accounted for by biological age, intelligence or inadequate schooling (WHO, 2011). It is hypothesized that DD relies, at least in part, on the impaired ability to shift attention to relevant information (Hari and Renvall, 2001). On a neurophysiological level, attention is represented by alpha oscillations (about 8–12 Hz). In adults with DD, recent studies showed altered alpha oscillations during the processing of auditory and visual stimuli (Dhar et al., 2010) but also in resting state (Papagiannopoulou and Lagopoulos, 2016). However, until today, there is only sparse knowledge on auditory attention and the underlying neural mechanisms in adolescents with DD. Methods A sample of adolescents with diagnosed DD (Mean age: 14.29 years, SD: 1.46) and an age-matched control group (CG) participated in this study. In balanced order they performed two auditory spatial-cueing paradigms. In the exogenous paradigm, cue and target tones were presented monaurally at Stimulus onset Asynchrony (SOA) 100 ms and 250 ms. 50% of the trials consisted of valid cues. Catch trials (cue only) were intermittently presented. Participants had to respond as fast as possible whenever a target tone was presented. In the endogenous paradigm, cue and target tones were presented at SOAs between 1000 ms and 1500 ms. 80% of the trials consisted of valid cues. During both tasks EEG data was continuously recorded. Results While the CG showed an inhibition of return (IOR) i.e. reduced RT at invalid compared to valid cues with SOA 250 ms, no such effect was present in the DD sample. ERPs measured at the vertex electrode revealed prolonged peak latencies for target stimuli for both valid and invalid trials at SOA 250 ms in DD compared to CG. In the endogenous task, we found increased alpha desynchronization in the post-cue phase over left auditory cortex (AC) regions in DD compared to CG. Notably, this pattern was found irrespective of the site at which the cue was presented. Discussion The lack of an IOR in adolescents with DD demonstrates a decreased withdrawal of attention and suggests reduced orienting towards novel locations. Considering the endogenous task, our findings indicate a maladaptive inhibition of the left AC in DD that, in turn, might account for dysfunctional processing of incoming acoustic information. Our findings suggest altered auditory attention as potential basis for higher order language skills. Future studies investigating attention processes in pre-reading children will shed more light on whether attention is a cause of impaired language skills rather than the consequence of a maladaptive compensation over the course of reading/ writing acquisition.

Background Healthy aging is typically associated with impairment in various cognitive abilities such as memory, selective attention or executive functions. Less well observed is the fact that also language functions in general and speech processing in particular seems to be affected by age. This impairment is partly caused by pathologies of the peripheral auditory nervous system and central auditory decline and in some part also by a cognitive decay. Aims This cross-sectional electroencephalography (EEG) study investigates temporally early electrophysiological correlates of auditory related selective attention in young (20–32 years) and older (60–74 years) healthy adults. Material and methods In two independent tasks, we systematically modulate the subjects' focus of attention by presenting words and pseudowords as targets and white noise stimuli as distractors. Results Behavioral data showed no difference in task accuracy between the two age samples irrespective of the modulation of attention. However, our work is the first to show that the N1- and the P2 component evoked by speech and nonspeech stimuli are specifically modulated in older adults and young adults depending on the subjects' focus of attention. Conclusion This finding is particularly interesting in that the age-related differences in AEPs may be reflecting levels of processing that are not mirrored by the behavioral measurements.

Transcranial electrical stimulation (tES) has become a valuable research tool for the investigation of neurophysiological processes underlying human action and cognition. In recent years, striking evidence for the neuromodulatory effects of transcranial direct current stimulation (tDCS), transcranial alternating current stimulation (tACS), and transcranial random noise stimulation (tRNS) has emerged. However, while the wealth of knowledge has been gained about tES in the motor domain and, to a lesser extent, about its ability to modulate human cognition, surprisingly little is known about its impact on perceptual processing, particularly in the auditory domain. Moreover, while only a few studies systematically investigated the impact of auditory tES, it has already been applied in a large number of clinical trials, leading to a remarkable imbalance between basic and clinical research on auditory tES. Here, we review the state of the art of tES application in the auditory domain focussing on the impact of neuromodulation on acoustic perception and its potential for clinical application in the treatment of auditory related disorders.

Background Normal aging is accompanied by a functional decline in processing temporal features of spoken language, such as voice onset time (VOT). On an electrophysiological level, this finding is paralleled by altered patterns of gamma oscillations. Objective Using 40 Hz transcranial alternating current stimulation (tACS) over the bilateral auditory cortex, this study aims to compare the effect of tACS to modulate VOT-processing in samples of healthy young and older adults. Methods Twenty-five healthy young (age 20–35 years) and 20 older adults (age 60–75 years) participated in this study. Presented with an auditory phoneme categorization task participants received 40 Hz and 6 Hz tACS on two consecutive sessions. Results While 40 Hz tACS diminished task accuracy in young adults, older adults benefitted from this stimulation resulting in a more precise phonetic categorization. Conclusion The results of the study are discussed with respect to the non-linear relationship between gamma oscillations in the vicinity of the auditory cortex and VOT-processing. The present findings are promising in the context of an intervention for subjects with impaired ability to process temporal acoustic features in the speech signal.

Theoretical background Decoding acoustic speech signals requires to parse the acoustic stream into linguistically meaningful units, and to extract stress and prosody. Accordingly, the temporal and spectral resolution of the auditory cortex is essential in successful speech processing ( Rosen, 1992 ). The reduced acuity in discriminating these features is considered as key factor in dyslexia ( Goswami, 2011 ) but is also a consequence of normal aging ( Walton, 2010 ). Transcranial random noise stimulation (tRNS), a non-invasive brain stimulation technique that applies weak currents at multiple frequencies, is assumed to modulate the excitability of neural assemblies and, in consequence, the reactivity of cortical structures. However, the effect of tRNS on the auditory system is yet unclear. The present study aimed to fill this gap by investigating the impact of tRNS on basic auditory feature processing. Considering the hemispheric specialization of the auditory system in analyzing theses features we hypothesized that a site-specific tRNS application leads to domain-specific changes in auditory features processing. In concrete, tRNS over the left auditory cortex (lAC) was hypothesized to affect the temporal domain while tRNS over the right auditory cortex (rAC) was assumed to modulate the spectral domain. Finally, we investigated whether tRNS over bilateral auditory cortex (bAC) leads to a modulation of both dimensions. Methods On four days 18 healthy, right-handed adults received 30 min. tRNS at an intensity of 1.5 mA either over the lAC, over the rAC, over bAC or sham stimulation while continuous EEG was recorded. The task consisted of the detection of a deviant tone in a sequence of three sine wave tones. While the duration of all tones and the pitch of the standard tones was kept stable, the inter-tone interval (temporal information) and the frequency in that the deviant tone differed from the standard tones (spectral information) was modulated. Results In result, we found that tRNS applied over the lAC and over bAC inferred with task performance. This effect was specifically modulated by the inter-tone interval between the tone triplets but not by the frequency of the deviant tone. On the electrophysiological level, this finding was paralleled by feature-specific brain response patterns in the alpha and the lower gamma band. Discussion Our findings demonstrate the functional relevance of the lAC in processing temporal features. Most importantly, we proved tRNS as a valid technique to modulate basic auditory perception. The current results stress the need for further investigations on the effect of tRNS in the auditory domain. Systematically assessing the most effective stimulation parameters provides the basis for the clinical application of tRNS in the context of speech and language disorders.

Introduction The impaired processing of rapidly changing auditory temporal features has been suggested as one key factor in development dyslexia. Voice onset time (VOT), a short period between the release of closure and the start of voicing enables to distinguish categorically between voiced (i.e./da/) and voiceless stop-consonants (i.e./ta/). VOT thus requires the precise perception and categorization of rapidly changing temporal information in the speech signal and can be considered as an important acoustic cue for the encoding of linguistically relevant information. Recently, non-invasive transcranial electrical stimulation has been suggested as a valid tool to modulate VOT-processing in healthy subjects (Heimrath et al., 2015; Rufener et al., 2016). Objectives The current study strives to assess the potential of transcranial alternating current stimulation (tACS) and transcranial random noise stimulation (tRNS) on VOT-categorization in dyslexic participants. Material and methods 16 adult dyslexics (age 20–50 years) with normal hearing acuity participated in this study. All participants received on three consecutive sessions either 20 min of tRNS (100–640 Hz), 40 Hz-tACS or sham stimulation over bilateral auditory cortex region while they performed an auditory phoneme categorization task. Subsequently, a passive MMN-paradigm with a standard stimulus (voiced/da/, VOT 20 ms), two within-category deviants (VOT 24 ms, VOT 28 ms) and to between-category deviants (voiceless/ta/, VOT 32 ms, VOT 36 ms) was performed. EEG-data were recorded from three midline electrodes (Fz, Cz, Pz) with the reference electrode at the tip of the nose. Results We found enhanced MMN-amplitudes at the Fz-electrode for both within-category deviants after 40 Hz-tACS compared to tRNS and sham. This finding indicates that tACS but not tRNS improves the auditory temporal resolution in patients with dyslexia. Conclusion Our data provides first evidence for a superior effect of tACS compared to tRNS on VOT-processing in dyslexic participants. Of note, we found increased temporal precision of the auditory system after a single session of VOT-categorization accompanied by tACS. Our finding emphasizes the clinical application of tACS in the context of impaired speech processing.

This EEG-study aims to investigate age-related differences in the neural oscillation patterns during the processing of temporally modulated speech. Viewing from a lifespan perspective, we recorded the electroencephalogram (EEG) data of three age samples: young adults, middle-aged adults and older adults. Stimuli consisted of temporally degraded sentences in Swedish-a language unfamiliar to all participants. We found age-related differences in phonetic pattern matching when participants were presented with envelope-degraded sentences, whereas no such age-effect was observed in the processing of fine-structure-degraded sentences. Irrespective of age, during speech processing the EEG data revealed a relationship between envelope information and the theta band (4-8 Hz) activity. Additionally, an association between fine-structure information and the gamma band (30-48 Hz) activity was found. No interaction, however, was found between acoustic manipulation of stimuli and age. Importantly, our main finding was paralleled by an overall enhanced power in older adults in high frequencies (gamma: 30-48 Hz). This occurred irrespective of condition. For the most part, this result is in line with the Asymmetric Sampling in Time framework (Poeppel in Speech Commun 41:245-255, 2003), which assumes an isomorphic correspondence between frequency modulations in neurophysiological patterns and acoustic oscillations in spoken language. We conclude that speech-specific neural networks show strong stability over adulthood, despite initial processes of cortical degeneration indicated by enhanced gamma power. The results of our study therefore confirm the concept that sensory and cognitive processes undergo multidirectional trajectories within the context of healthy aging.