Nadia Müller

Top-down modulation of the auditory Steady State Response in a task-switch paradigm

Auditory selective attention is an important mechanism for top-down selection of the vast amount of auditory information our perceptual system is exposed to. In the present study, the impact of attention on auditory steady-state responses is investigated. This issue is still a matter of debate and recent findings point to a complex pattern of attentional effects on the auditory steady state response (aSSR). The present study aimed at shedding light on the involvement of ipsilateral and contralateral activations to the attended sound taking into account hemispheric differences and a possible dependency on modulation frequency. In aid of this, a dichotic listening experiment was designed using amplitude-modulated tones that were presented to the left and right ear simultaneously. Participants had to detect target tones in a cued ear while their brain activity was assessed using MEG. Thereby, a modulation of the aSSR by attention could be revealed, interestingly restricted to the left hemisphere and 20 Hz responses: contralateral activations were enhanced while ipsilateral activations turned out to be reduced. Thus, our findings support and extend recent findings, showing that auditory attention can influence the aSSR, but only under specific circumstances and in a complex pattern regarding the different effects for ipsilateral and contralateral activations.

Alpha rhythms in audition: Cognitive and clinical perspectives

Like the visual and the sensorimotor systems, the auditory system exhibits pronounced alpha-like resting oscillatory activity. Due to the relatively small spatial extent of auditory cortical areas, this rhythmic activity is less obvious and frequently masked by non-auditory alpha-generators when recording non-invasively using magnetoencephalography (MEG) or electroencephalography (EEG). Following stimulation with sounds, marked desynchronizations can be observed between 6 and 12 Hz, which can be localized to the auditory cortex. However knowledge about the functional relevance of the auditory alpha rhythm has remained scarce so far. Results from the visual and sensorimotor system have fuelled the hypothesis of alpha activity reflecting a state of functional inhibition. The current article pursues several intentions: (1) Firstly we review and present own evidence (MEG, EEG, sEEG) for the existence of an auditory alpha-like rhythm independent of visual or motor generators, something that is occasionally met with skepticism. (2) In a second part we will discuss tinnitus and how this audiological symptom may relate to reduced background alpha. The clinical part will give an introduction into a method which aims to modulate neurophysiological activity hypothesized to underlie this distressing disorder. Using neurofeedback, one is able to directly target relevant oscillatory activity. Preliminary data point to a high potential of this approach for treating tinnitus. (3) Finally, in a cognitive neuroscientific part we will show that auditory alpha is modulated by anticipation/expectations with and without auditory stimulation. We will also introduce ideas and initial evidence that alpha oscillations are involved in the most complex capability of the auditory system, namely speech perception. The evidence presented in this article corroborates findings from other modalities, indicating that alpha-like activity functionally has an universal inhibitory role across sensory modalities.

On the Variability of the McGurk Effect: Audiovisual Integration Depends on Prestimulus Brain States

The McGurk effect demonstrates the influence of visual cues on auditory perception. Mismatching information from both sensory modalities can fuse to a novel percept that matches neither the auditory nor the visual stimulus. This illusion is reported in 60-80% of trials. We were interested in the impact of ongoing brain oscillations-indexed by fluctuating local excitability and interareal synchronization-on upcoming perception of identical stimuli. The perception of the McGurk effect is preceded by high beta activity in parietal, frontal, and temporal areas. Beta activity is pronounced in the left superior temporal gyrus (lSTG), which is considered as a site of multimodal integration. This area is functionally (de)coupled to distributed frontal and temporal regions in illusion trials. The disposition to fuse multisensory information is enhanced as the lSTG is more strongly coupled to frontoparietal regions. Illusory perception is accompanied by a decrease in poststimulus theta-band activity in the cuneus, precuneus, and left superior frontal gyrus. Event-related activity in the left middle temporal gyrus is pronounced during illusory perception. Thus, the McGurk effect depends on fluctuating brain states suggesting that functional connectedness of left STS at a prestimulus stage is crucial for an audiovisual percept.

Lateralized Auditory Cortical Alpha Band Activity and Interregional Connectivity Pattern Reflect Anticipation of Target Sounds

The anticipation of stimuli facilitates the top-down preparation of neuronal tissue involved in the processing of forthcoming targets. Increasing evidence in the visual modality emphasizes the anticipatory adjustment of visual cortex excitability through modulations of oscillatory alpha power. In the auditory system, however, this relationship has not yet been established. Furthermore, the association between anticipatory modulations of auditory alpha power and a potential top-down network within these anticipatory preparation processes remains unexplained. To disclose these processes, we recorded magnetoencephalography while visually cuing participants to attend to either ear and to anticipate forthcoming auditory stimuli. For the cue-stimulus phase, we expected an asymmetric modulation of auditory alpha power when attending to the left or right ear, assuming that frontoparietal regions would phase synchronize with the auditory cortex in an asymmetric pattern. Beamformer source solutions demonstrate an asymmetric modulation of auditory alpha power following visual cues expressed in a strong right auditory alpha power increase when attending to the right ear. Furthermore, the right auditory cortex is functionally connected to the frontal eye fields during the ipsilateral alpha increase. Altogether, these results contribute significantly to the understanding of how auditory anticipation acts on a local as well as on a network level.

Rapid increases of gamma power in the auditory cortex following noise trauma in humans

Tinnitus is an auditory perception in the absence of any external sound source. It has been suggested that tinnitus is related to enhanced synchronization of neuronal activity in the auditory cortex. Usually a hearing damage can be identified suggesting auditory deprivation to central auditory regions to be fundamental for neurophysiological processes related to tinnitus. Until now, human research has been conducted on patients with chronic tinnitus (> 6 months). However, neuronal activity accompanying auditory deprivation and putatively tinnitus may not remain constant over time, making it difficult to directly relate outcomes of current animal studies (acute tinnitus) to chronic tinnitus in humans, and vice versa. We investigated 14 amateur rock musicians who frequently reported a short‐term tinnitus immediately after band practice. Magnetoencephalographic resting‐state recordings, audiometry and tinnitus testing were performed at two separate occasions: with and without previous exposure to loud music. Analyses revealed that transient tinnitus was accompanied by temporary hearing loss in both ears and increased gamma activity in the right auditory cortex in 13 out of 14 cases. Additionally, tinnitus frequency was strongly correlated to hearing loss. Analogous to animal studies, our results show for the first time in humans that noise trauma leads rapidly to increased neuronal synchrony in the auditory cortex. Importantly, the strongly right‐lateralized effect implies that it does not reflect tinnitus percept per se. This could rather have been triggered by greater discontinuities of hearing loss at high frequencies that were particularly pronounced in the left ear.

Loss of alpha power is related to increased gamma synchronization—A marker of reduced inhibition in tinnitus?

Tinnitus is the perception of sound in the absence of any external auditory stimulus. Based on previous research we have proposed a framework which postulates that the reduction of ongoing inhibitory alpha activity in tinnitus subjects favors a synchronization of neurons in the gamma frequency range while in a resting state. In the present work we are validating the existence of an inverse relationship between auditory gamma and alpha activity in tinnitus and control subjects using Magnetoencephalography. Tinnitus subjects exhibited a significantly steeper slope of the regression line compared to controls, presumably because a greater number of subjects concurrently exhibited low alpha and high gamma power. Therefore, the role of the alpha-gamma pattern is discussed regarding its possible implication for the generation of tinnitus.

Now I am Ready—Now I am not: The Influence of Pre-TMS Oscillations and Corticomuscular Coherence on Motor-Evoked Potentials

There is a growing body of research on the functional role of oscillatory brain activity. However, its relation to functional connectivity has remained largely obscure. In the sensorimotor system, movement-related changes emerge in the α (8-14 Hz) and β (15-30 Hz) range (event-related desynchronization, ERD, before and during movement; event-related synchronization, ERS, after movement offset). Some studies suggest that β-ERS may functionally inhibit new movements. According to the gating-by-inhibition framework ( Jensen and Mazaheri 2010), we expected that the ERD would go along with increased corticomuscular coupling, and vice versa. By combining transcranial magnetic stimulation (TMS) and electroencephalography, we were directly able to test this hypothesis. In a reaction time task, single TMS pulses were delivered randomly during ERD/ERS to the motor cortex. The motor-evoked potential (MEP) was then related to the β and α frequencies and corticomuscular coherence. Results indicate that MEPs are smaller when preceded by high pre-TMS β-band power and low pre-TMS α-band corticomuscular coherence (and vice versa) in a network of motor-relevant areas comprising frontal, parietal, and motor cortices. This confirms that an increase in rhythms that putatively reflect functionally inhibited states goes along with weaker coupling of the respective brain regions.

Short-term effects of single repetitive TMS sessions on auditory evoked activity in patients with chronic tinnitus.

Subjective tinnitus is the perception of a sound without any external source. Repetitive transcranial magnetic stimulation (rTMS) has been examined as a treatment tool for chronic tinnitus for several years trying to target hyperactivity/abnormal synchronization within the auditory cortex putatively underlying the auditory phantom percept. However, its exact impact on auditory cortical activity remains largely unknown. This studys objective was to systematically examine changes in auditory responses (N1, auditory steady-state response [aSSR]) measured by means of magnetoencephalography after single sessions of stimulation with different TMS paradigms. Subjects with chronic tinnitus (n = 10) underwent five sessions of rTMS in which they received one of five different stimulation protocols (1 Hz, individual alpha frequency, continuous theta burst stimulation [cTBS], intermittent theta burst stimulation [iTBS], and sham) in randomized order using a single-blind study design. Cortical steady-state responses to 40 Hz amplitude-modulated tones were measured before and after each magnetic stimulation protocol. The results demonstrate a reduction of the cortical response to the auditory steady-state stimulus after magnetic stimulation, whereas the N1 response was slightly enhanced or remained unchanged. Furthermore, reduction of the aSSR was driven by effects of iTBS, cTBS, and 1 Hz stimulation. Correspondingly, behavioral measures demonstrated the greatest reduction of tinnitus loudness after the respective rTMS protocols. The current study offers an interesting insight into the effects of rTMS on auditory cortical activity. The results of the study are discussed in the context of current limitations of TMS for the treatment of chronic tinnitus.

Prestimulus Beta Power and Phase Synchrony Influence the Sound-Induced Flash Illusion

We investigated the sound-induced flash illusion, an example for the influence of auditory information on visual perception. It consists of the perception of 2 visual stimuli upon the presentation of a single visual stimulus alongside 2 auditory stimuli. We used magnetoencephalography to assess the influence of prestimulus oscillatory activity on varying the perception of invariant stimuli. We compared cortical activity from trials in which subjects perceived an illusion with trials in which subjects did not perceive the illusion, keeping the stimulation fixed. Subjects perceived the illusion in approximately 50% of trials. Prior to the illusion, we found stronger beta-band power in left temporal sensors, localized to the left middle temporal gyrus. Illusory perceptions were preceded by increased beta-band phase synchrony between the left middle temporal gyrus and auditory areas as well as by decreased phase synchrony with visual areas. Alpha-band phase synchrony between visual and temporal, parietal, and frontal cortical areas as well as alpha-band phase synchrony between auditory and visual areas were modulated. This supports and extends reports on the influence of brain states prior to stimulation on subsequent perception. We suggest that prestimulus local and network activities form predispositions if sensory streams will be integrated.

Oscillatory Alpha Modulations in Right Auditory Regions Reflect the Validity of Acoustic Cues in an Auditory Spatial Attention Task

Anticipation of targets in the left or right hemifield leads to alpha modulations in posterior brain areas. Recently using magnetoencephalography, we showed increased right auditory alpha activity when attention was cued ipsilaterally. Here, we investigated the issue how cue validity itself influences oscillatory alpha activity. Acoustic cues were presented either to the right or left ear, followed by a compound dichotically presented target plus distractor. The preceding cue was either informative (75% validity) or uninformative (50%) about the location of the upcoming target. Cue validity × side-related alpha modulations were identified in pre- and posttarget periods in a right lateralized network, comprising auditory and nonauditory regions. This replicates and extends our previous finding of the right hemispheric dominance of auditory attentional modulations. Importantly, effective connectivity analysis showed that, in the pretarget period, this effect is accompanied by a pronounced and time-varying connectivity pattern of the right auditory cortex to the right intraparietal sulcus (IPS), with influence of IPS on superior temporal gyrus dominating at earlier intervals of the cue-target period. Our study underlines the assumption that alpha oscillations may play a similar functional role in auditory cortical regions as reported in other sensory modalities and suggests that these effects may be mediated via IPS.