Modulations of neural activity in auditory streaming caused by spectral and temporal alternation in subsequent stimuli: a magnetoencephalographic study

Background: The aim of the present study was to identify a specific neuronal correlate underlying the preattentive auditory stream segregation of subsequent sound patterns alternating in spectral or temporal cues. Fifteen participants with normal hearing were presented with series’ of two consecutive ABA auditory tone-triplet sequences, the initial triplets being the Adaptation sequence and the subsequent triplets being the Test sequence. In the first experiment, the frequency separation (delta-f) between A and B tones in the sequences was varied by 2, 4 and 10 semitones. In the second experiment, a constant delta-f of 6 semitones was maintained but the Inter-Stimulus Intervals (ISIs) between A and B tones were varied. Auditory evoked magnetic fields (AEFs) were recorded using magnetoencephalography (MEG). Participants watched a muted video of their choice and ignored the auditory stimuli. In a subsequent behavioral study both MEG experiments were replicated to provide information about the participants’ perceptual state. Results: MEG measurements showed a significant increase in the amplitude of the B-tone related P1 component of the AEFs as delta-f increased. This effect was seen predominantly in the left hemisphere. A significant increase in the amplitude of the N1 component was only obtained for a Test sequence delta-f of 10 semitones with a prior Adaptation sequence of 2 semitones. This effect was more pronounced in the right hemisphere. The additional behavioral data indicated an increased probability of two-stream perception for delta-f = 4 and delta-f = 10 semitones with a preceding Adaptation sequence of 2 semitones. However, neither the neural activity nor the perception of the successive streaming sequences were modulated when the ISIs were alternated. Conclusions: Our MEG experiment demonstrated differences in the behavior of P1 and N1 components during the automatic segregation of sounds when induced by an initial Adaptation sequence. The P1 component appeared enhanced in all Test-conditions and thus demonstrates the preceding context effect, whereas N1 was specifically modulated only by large delta-f Test sequences induced by a preceding small delta-f Adaptation sequence. These results suggest that P1 and N1 components represent at least partially-different systems that underlie the neural representation of auditory streaming

Perceptual organization of auditory streaming-task relies on neural entrainment of the stimulus-presentation rate: MEG evidence

Background: Humans are able to extract regularities from complex auditory scenes in order to form perceptually meaningful elements. It has been shown previously that this process depends critically on both the temporal integration of the sensory input over time and the degree of frequency separation between concurrent sound sources. Our goal was to examine the relationship between these two aspects by means of magnetoencephalography (MEG). To achieve this aim, we combined time-frequency analysis on a sensor space level with source analysis. Our paradigm consisted of asymmetric ABA-tone triplets wherein the B-tones were presented temporally closer to the first A-tones, providing different tempi within the same sequence. Participants attended to the slowest B-rhythm whilst the frequency separation between tones was manipulated (0-, 2-, 4- and 10-semitones). Results: The results revealed that the asymmetric ABA-triplets spontaneously elicited periodic-sustained responses corresponding to the temporal distribution of the A-B and B-A tone intervals in all conditions. Moreover, when attending to the B-tones, the neural representations of the A- and B-streams were both detectable in the scenarios which allow perceptual streaming (2-, 4- and 10-semitones). Alongside this, the steady-state responses tuned to the presentation of the B-tones enhanced significantly with increase of the frequency separation between tones. However, the strength of the B-tones related steady-state responses dominated the strength of the A-tones responses in the 10-semitones condition. Conversely, the representation of the A-tones dominated the B-tones in the cases of 2- and 4-semitones conditions, in which a greater effort was required for completing the task. Additionally, the P1 evoked fields’ component following the B-tones increased in magnitude with the increase of inter-tonal frequency difference. Conclusions: The enhancement of the evoked fields in the source space, along with the B-tones related activity of the time-frequency results, likely reflect the selective enhancement of the attended B-stream. The results also suggested a dissimilar efficiency of the temporal integration of separate streams depending on the degree of frequency separation between the sounds. Overall, the present findings suggest that the neural effects of auditory streaming could be directly captured in the time-frequency spectrum at the sensor-space level.<br

A Large N400 but No BOLD Effect – Comparing Source Activations of Semantic Priming in Simultaneous EEG-fMRI

Numerous studies have reported neurophysiological effects of semantic priming in electroencephalography (EEG) and in functional magnetic resonance imaging (fMRI). Because of differing methodological constraints, the comparability of the observed effects remains unclear. To directly compare EEG and fMRI effects and neural sources of semantic priming, we conducted a semantic word-picture priming experiment while measuring EEG and fMRI simultaneously. The visually presented primes were pseudowords, words unrelated to the target, semantically related words and the identical names of the target. Distributed source analysis of the event-related potentials (ERPs) successfully revealed a large effect of semantic prime-target relatedness (the N400 effect), which was driven by activations in a left-temporal source region. However, no significantly differing activations between priming conditions were found in the fMRI data. Our results support the notion that, for joint interpretations of existing EEG and fMRI studies of semantic priming, we need to fully appreciate the respective methodological limitations. Second, they show that simultaneous EEG-fMRI, including ERP source localization, is a feasible and promising methodological advancement for the investigation of higher-cognitive processes. Third, they substantiate the finding that, compared to fMRI, ERPs are often more sensitive to subtle cognitive effects

Playing and Listening to Tailor-Made Notched Music: Cortical Plasticity Induced by Unimodal and Multimodal Training in Tinnitus Patients

Background. The generation and maintenance of tinnitus are assumed to be based on maladaptive functional cortical reorganization. Listening to modified music, which contains no energy in the range of the individual tinnitus frequency, can inhibit the corresponding neuronal activity in the auditory cortex. Music making has been shown to be a powerful stimulator for brain plasticity, inducing changes in multiple sensory systems. Using magnetoencephalographic (MEG) and behavioral measurements we evaluated the cortical plasticity effects of two months of (a) active listening to (unisensory) versus (b) learning to play (multisensory) tailor-made notched music in nonmusician tinnitus patients. Taking into account the fact that uni- and multisensory trainings induce different patterns of cortical plasticity we hypothesized that these two protocols will have different affects. Results. Only the active listening (unisensory) group showed significant reduction of tinnitus related activity of the middle temporal cortex and an increase in the activity of a tinnitus-coping related posterior parietal area. Conclusions. These findings indicate that active listening to tailor-made notched music induces greater neuroplastic changes in the maladaptively reorganized cortical network of tinnitus patients while additional integration of other sensory modalities during training reduces these neuroplastic effects

Enhancing Inhibition-Induced Plasticity in Tinnitus – Spectral Energy Contrasts in Tailor-Made Notched Music Matter

Chronic tinnitus seems to be caused by reduced inhibition among frequency selective neurons in the auditory cortex. One possibility to reduce tinnitus perception is to induce inhibition onto over-activated neurons representing the tinnitus frequency via tailor-made notched music (TMNM). Since lateral inhibition is modifiable by spectral energy contrasts, the question arises if the effects of inhibition-induced plasticity can be enhanced by introducing increased spectral energy contrasts (ISEC) in TMNM. Eighteen participants suffering from chronic tonal tinnitus, pseudo randomly assigned to either a classical TMNM or an ISEC-TMNM group, listened to notched music for three hours on three consecutive days. The music was filtered for both groups by introducing a notch filter centered at the individual tinnitus frequency. For the ISEC-TMNM group a frequency bandwidth of 3/8 octaves on each side of the notch was amplified, additionally, by about 20 dB. Before and after each music exposure, participants rated their subjectively perceived tinnitus loudness on a visual analog scale. During the magnetoencephalographic recordings, participants were stimulated with either a reference tone of 500 Hz or a test tone with a carrier frequency representing the individual tinnitus pitch. Perceived tinnitus loudness was significantly reduced after TMNM exposure, though TMNM type did not influence the loudness ratings. Tinnitus related neural activity in the N1m time window and in the so called tinnitus network comprising temporal, parietal and frontal regions was reduced after TMNM exposure. The ISEC-TMNM group revealed even enhanced inhibition-induced plasticity in a temporal and a frontal cortical area. Overall, inhibition of tinnitus related neural activity could be strengthened in people affected with tinnitus by increasing spectral energy contrast in TMNM, confirming the concepts of inhibition-induced plasticity via TMNM and spectral energy contrasts

Functional oropharyngeal sensory disruption interferes with the cortical control of swallowing

<p>Abstract</p> <p>Background</p> <p>Sensory input is crucial to the initiation and modulation of swallowing. From a clinical point of view, oropharyngeal sensory deficits have been shown to be an important cause of dysphagia and aspiration in stroke patients. In the present study we therefore investigated effects of functional oropharyngeal disruption on the cortical control of swallowing. We employed whole-head MEG to study cortical activity during self-paced volitional swallowing with and without topical oropharyngeal anesthesia in ten healthy subjects. A simple swallowing screening-test confirmed that anesthesia caused swallowing difficulties with decreased swallowing speed and reduced volume per swallow in all subjects investigated. Data were analyzed by means of synthetic aperture magnetometry (SAM) and the group analysis of the individual SAM data was performed using a permutation test.</p> <p>Results</p> <p>The analysis of normal swallowing revealed bilateral activation of the mid-lateral primary sensorimotor cortex. Oropharyngeal anesthesia led to a pronounced decrease of both sensory and motor activation.</p> <p>Conclusion</p> <p>Our results suggest that a short-term decrease in oropharyngeal sensory input impedes the cortical control of swallowing. Apart from diminished sensory activity, a reduced activation of the primary motor cortex was found. These findings facilitate our understanding of the pathophysiology of dysphagia.</p

Targeting Heterogeneous Findings in Neuronal Oscillations in Tinnitus: Analyzing MEG Novices and Mental Health Comorbidities

Tinnitus is a prevalent phenomenon and bothersome for people affected by it. Its occurrence and maintenance have a clear neuroscientific tie and one aspect are differences in the neuronal oscillatory pattern, especially in auditory cortical areas. As studies in this field come to different results, the aim of this study was to analyze a large number of participants to achieve more stable results. Furthermore, we expanded our analysis to two variables of potential influence, namely being a novice to neuroscientific measurements and the exclusion of psychological comorbidities. Oscillatory brain activity of 88 subjects (46 with a chronic tinnitus percept, 42 without) measured in resting state by MEG was investigated. In the analysis based on the whole group, in sensor space increased activity in the delta frequency band was found in tinnitus patients. Analyzing the subgroup of novices, a significant difference in the theta band emerged additionally to the delta band difference (sensor space). Localizing the origin of the activity, we found a difference in theta and gamma band for the auditory regions for the whole group and the same significant difference in the subgroup of novices. However, no differences in oscillatory activity were observed between tinnitus and control groups once subjects with mental health comorbidity were excluded. Against the background of previous studies, the study at hand underlines the fragility of the results in the field of neuronal cortical oscillations in tinnitus. It supports the body of research arguing for low frequency oscillations and gamma band activity as markers associated with tinnitus

Maladaptive alterations of resting state cortical network in Tinnitus: A directed functional connectivity analysis of a larger MEG data set

The present study used resting state MEG whole-head recordings to identify how chronic tonal tinnitus relates to altered functional connectivity of brain’s intrinsic cortical networks. Resting state MEG activity of 40 chronic tinnitus patients and 40 matched human controls was compared identifying significant alterations in intrinsic networks of the tinnitus population. Directed functional connectivity of the resting brain, at a whole cortex level, was estimated by means of a statistical comparison of the estimated phase Transfer Entropy (pTE) between the time-series of cortical activations, as reconstructed by LORETA. As pTE identifies the direction of the information flow, a detailed analysis of the connectivity differences between tinnitus patients and controls was possible. Results indicate that the group of tinnitus patients show increased connectivity from right dorsal prefrontal to right medial temporal areas. Our results go beyond previous findings by indicating that the role of the left para-hippocampal area is dictated by a modulation from dmPFC; a region that is part of the dorsal attention network (DAN), as well as implicated in the regulation of emotional processing. Additionally, this whole cortex analysis showed a crucial role of the left inferior parietal cortex, which modulated the activity of the right superior temporal gyrus, providing new hypotheses for the role of this area within the context of current tinnitus models. Overall, these maladaptive alterations of the structure of intrinsic cortical networks show a decrease in efficiency and small worldness of the resting state network of tinnitus patients, which is correlated to tinnitus distress

Study Protocol: Münster Tinnitus randomized controlled Clinical Trial-2013 based on tailor-made notched music treatment (TMNMT)

Background: Tinnitus is a result of hyper-activity/hyper-synchrony of auditory neurons coding the tinnitus frequency, which has developed to synchronous mass activity owing the lack of inhibition. We assume that removal of exactly these frequency components from an auditory stimulus will cause the brain to reorganize around tonotopic regions coding the tinnitus frequency. Based on this assumption a novel treatment for tonal tinnitus - tailor-made notched music training (TMNMT) (Proc Natl Acad Sci USA 107:1207–1210, 2010; Ann N Y Acad Sci 1252:253–258, 2012; Frontiers Syst Neurosci 6:50, 2012) has been introduced and will be tested in this clinical trial on a large number of tinnitus patients. Methods and design: A randomized controlled trial (RCT) in parallel group design will be performed in a double-blinded manner. The choice of the intervention we are going to apply is based on two “proof of concept” studies in humans (Proc Natl Acad Sci USA 107:1207–1210, 2010; Ann N Y Acad Sci 1252:253–258, 2012; Frontiers Syst Neurosci 6:50, 2012; PloS One 6(9):e24685, 2011) and on a recent animal study (Front Syst Neurosci 7:21, 2013). The RCT includes 100 participants with chronic, tonal tinnitus who listened to tailor-made notched music (TMNM) for two hours a day for three months. The effect of TMNMT is assessed by the tinnitus handicap questionnaire and visual analogue scales (VAS) measuring perceived tinnitus loudness, distress and handicap. Discussion: This is the first randomized controlled trial applying TMNMT on a larger number of patients with tonal tinnitus. Our data will verify more securely and reliably the effectiveness of this kind of completely non-invasive and low-cost treatment approach on tonal tinnitus

Auditory Categorization of Man-Made Sounds Versus Natural Sounds by Means of MEG Functional Brain Connectivity

Previous neuroimaging studies have shown that sounds can be discriminated due to living-related or man-made-related characteristics and involve different brain regions. However, these studies have mainly provided source space analyses, which offer simple maps of activated brain regions but do not explain how regions of a distributed system are functionally organized under a specific task. In the present study, we aimed to further examine the functional connectivity of the auditory processing pathway across different categories of non-speech sounds in healthy adults, by means of MEG. Our analyses demonstrated significant activation and interconnection differences between living and man-made object sounds, in the prefrontal areas, anterior-superior temporal gyrus (aSTG), posterior cingulate cortex (PCC), and supramarginal gyrus (SMG), occurring within 80–120 ms post-stimulus interval. Current findings replicated previous ones, in that other regions beyond the auditory cortex are involved during auditory processing. According to the functional connectivity analysis, differential brain networks across the categories exist, which proposes that sound category discrimination processing relies on distinct cortical networks, a notion that has been strongly argued in the literature also in relation to the visual system