Speech rhythms and multiplexed oscillatory sensory coding in the human brain

Cortical oscillations are likely candidates for segmentation and coding of continuous speech. Here, we monitored continuous speech processing with magnetoencephalography (MEG) to unravel the principles of speech segmentation and coding. We demonstrate that speech entrains the phase of low-frequency (delta, theta) and the amplitude of high-frequency (gamma) oscillations in the auditory cortex. Phase entrainment is stronger in the right and amplitude entrainment is stronger in the left auditory cortex. Furthermore, edges in the speech envelope phase reset auditory cortex oscillations thereby enhancing their entrainment to speech. This mechanism adapts to the changing physical features of the speech envelope and enables efficient, stimulus-specific speech sampling. Finally, we show that within the auditory cortex, coupling between delta, theta, and gamma oscillations increases following speech edges. Importantly, all couplings (i.e., brain-speech and also within the cortex) attenuate for backward-presented speech, suggesting top-down control. We conclude that segmentation and coding of speech relies on a nested hierarchy of entrained cortical oscillations

A new unifying account of the roles of neuronal entrainment

Rhythms are a fundamental and defining feature of neuronal activity in animals including humans. This rhythmic brain activity interacts in complex ways with rhythms in the internal and external environment through the phenomenon of ‘neuronal entrainment’, which is attracting increasing attention due to its suggested role in a multitude of sensory and cognitive processes. Some senses, such as touch and vision, sample the environment rhythmically, while others, like audition, are faced with mostly rhythmic inputs. Entrainment couples rhythmic brain activity to external and internal rhythmic events, serving fine-grained routing and modulation of external and internal signals across multiple spatial and temporal hierarchies. This interaction between a brain and its environment can be experimentally investigated and even modified by rhythmic sensory stimuli or invasive and non-invasive neuromodulation techniques. We provide a comprehensive overview of the topic and propose a theoretical framework of how neuronal entrainment dynamically structures information from incoming neuronal, bodily and environmental sources. We discuss the different types of neuronal entrainment, the conceptual advances in the field, and converging evidence for general principles

Top-down control of alpha phase adjustment in anticipation of predictable visual stimuli

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Who is that? Brain networks and mechanisms for identifying individuals

Social animals can identify conspecifics by many forms of sensory input. However, whether the neuronal computations that support this ability to identify individuals rely on modality-independent convergence or involve ongoing synergistic interactions along the multiple sensory streams remains controversial. Direct neuronal measurements at relevant brain sites could address such questions, but this requires better bridging the work in humans and animal models. Here, we overview recent studies in nonhuman primates on voice and face identity-sensitive pathways and evaluate the correspondences to relevant findings in humans. This synthesis provides insights into converging sensory streams in the primate anterior temporal lobe (ATL) for identity processing. Furthermore, we advance a model and suggest how alternative neuronal mechanisms could be tested

Gamma oscillations in human primary somatosensory cortex reflect pain perception

Pain is a highly subjective sensation of inherent behavioral importance and is therefore expected to receive enhanced processing in relevant brain regions. We show that painful stimuli induce high-frequency oscillations in the electrical activity of the human primary somatosensory cortex. Amplitudes of these pain-induced gamma oscillations were more closely related to the subjective perception of pain than to the objective stimulus attributes. They correlated with participants' ratings of pain and were stronger for laser stimuli that caused pain, compared with the same stimuli when no pain was perceived. These findings indicate that gamma oscillations may represent an important mechanism for processing behaviorally relevant sensory information

On the role of neuronal oscillations in auditory cortical processing

Although it has been over 100 years since William James stated that everyone knows what attention is , its underlying neural mechanisms are still being debated today. The goal of this research was to describe the physiological mechanisms of auditory attention using direct electrophysiological recordings in macaque primary auditory cortex (A1). A major focus of my research was on the role ongoing neuronal oscillations play in attentional modulation of auditory responses in A1. For all studies, laminar profiles of synaptic activity, (indexed by current source density analysis) and concomitant firing patterns in local neurons (multiunit activity) were acquired simultaneously via linear array multielectrodes positioned in A1. The initial study of this dissertation examined the contribution of ongoing oscillatory activity to excitatory and inhibitory responses in A1 in passive (no task) conditions. Next, the function of ongoing oscillations in modulating the frequency tuning of A1 during an intermodal selective attention oddball task was investigated. The last study was aimed at establishing whether there is a hemispheric asymmetry in the way neuronal oscillations are utilized by attention, corresponding to that noted in humans. The results of the first study indicate that in passive conditions, ongoing oscillations reset by stimulus related inputs modulate both excitatory and inhibitory components of local neuronal ensemble responses in A1. The second set of experiments demonstrates that this mechanism is utilized by attention to modulate and sharpen frequency tuning. Finally, we show that as in humans, there appears to be a specialization of left A1 for temporal processing, as signified by greater temporal precision of neuronal oscillatory alignment. Taken together these results underline the importance of neuronal oscillations in perceptual processes, and the validity of the macaque monkey as a model of human auditory processing

Neural oscillatory signatures of auditory and audiovisual illusions

Questions of the relationship between human perception and brain activity can be approached from different perspectives: in the first, the brain is mainly regarded as a recipient and processor of sensory data. The corresponding research objective is to establish mappings of neural activity patterns and external stimuli. Alternatively, the brain can be regarded as a self-organized dynamical system, whose constantly changing state affects how incoming sensory signals are processed and perceived. The research reported in this thesis can chiefly be located in the second framework, and investigates the relationship between oscillatory brain activity and the perception of ambiguous stimuli. Oscillations are here considered as a mechanism for the formation of transient neural assemblies, which allows efficient information transfer. While the relevance of activity in distinct frequency bands for auditory and audiovisual perception is well established, different functional architectures of sensory integration can be derived from the literature. This dissertation therefore aims to further clarify the role of oscillatory activity in the integration of sensory signals towards unified perceptual objects, using illusion paradigms as tools of study. In study 1, we investigate the role of low frequency power modulations and phase alignment in auditory object formation. We provide evidence that auditory restoration is associated with a power reduction, while the registration of an additional object is reflected by an increase in phase locking. In study 2, we analyze oscillatory power as a predictor of auditory influence on visual perception in the sound-induced flash illusion. We find that increased beta-/ gamma-band power over occipitotemporal electrodes shortly before stimulus onset predicts the illusion, suggesting a facilitation of processing in polymodal circuits. In study 3, we address the question of whether visual influence on auditory perception in the ventriloquist illusion is reflected in primary sensory or higher-order areas. We establish an association between reduced theta-band power in mediofrontal areas and the occurrence of illusion, which indicates a top-down influence on sensory decision-making. These findings broaden our understanding of the functional relevance of neural oscillations by showing that different processing modes, which are reflected in specific spatiotemporal activity patterns, operate in different instances of sensory integration.Fragen nach dem Zusammenhang zwischen menschlicher Wahrnehmung und Hirnaktivität können aus verschiedenen Perspektiven adressiert werden: in der einen wird das Gehirn hauptsächlich als Empfänger und Verarbeiter von sensorischen Daten angesehen. Das entsprechende Forschungsziel wäre eine Zuordnung von neuronalen Aktivitätsmustern zu externen Reizen. Dieser Sichtweise gegenüber steht ein Ansatz, der das Gehirn als selbstorganisiertes dynamisches System begreift, dessen sich ständig verändernder Zustand die Verarbeitung und Wahrnehmung von sensorischen Signalen beeinflusst. Die Arbeiten, die in dieser Dissertation zusammengefasst sind, können vor allem in der zweitgenannten Forschungsrichtung verortet werden, und untersuchen den Zusammenhang zwischen oszillatorischer Hirnaktivität und der Wahrnehmung von mehrdeutigen Stimuli. Oszillationen werden hier als ein Mechanismus für die Formation von transienten neuronalen Zusammenschlüssen angesehen, der effizienten Informationstransfer ermöglicht. Obwohl die Relevanz von Aktivität in verschiedenen Frequenzbändern für auditorische und audiovisuelle Wahrnehmung gut belegt ist, können verschiedene funktionelle Architekturen der sensorischen Integration aus der Literatur abgeleitet werden. Das Ziel dieser Dissertation ist deshalb eine Präzisierung der Rolle oszillatorischer Aktivität bei der Integration von sensorischen Signalen zu einheitlichen Wahrnehmungsobjekten mittels der Nutzung von Illusionsparadigmen. In der ersten Studie untersuchen wir die Rolle von Leistung und Phasenanpassung in niedrigen Frequenzbändern bei der Formation von auditorischen Objekten. Wir zeigen, dass die Wiederherstellung von Tönen mit einer Reduktion der Leistung zusammenhängt, während die Registrierung eines zusätzlichen Objekts durch einen erhöhten Phasenangleich widergespiegelt wird. In der zweiten Studie analysieren wir oszillatorische Leistung als Prädiktor von auditorischem Einfluss auf visuelle Wahrnehmung in der sound-induced flash illusion. Wir stellen fest, dass erhöhte Beta-/Gamma-Band Leistung über occipitotemporalen Elektroden kurz vor der Reizdarbietung das Auftreten der Illusion vorhersagt, was auf eine Begünstigung der Verarbeitung in polymodalen Arealen hinweist. In der dritten Studie widmen wir uns der Frage, ob ein visueller Einfluss auf auditorische Wahrnehmung in der ventriloquist illusion sich in primären sensorischen oder übergeordneten Arealen widerspiegelt. Wir weisen einen Zusammenhang von reduzierter Theta-Band Leistung in mediofrontalen Arealen und dem Auftreten der Illusion nach, was einen top-down Einfluss auf sensorische Entscheidungsprozesse anzeigt. Diese Befunde erweitern unser Verständnis der funktionellen Bedeutung neuronaler Oszillationen, indem sie aufzeigen, dass verschiedene Verarbeitungsmodi, die sich in spezifischen räumlich-zeitlichen Aktivitätsmustern spiegeln, in verschiedenen Phänomenen von sensorischer Integration wirksam sind

Perceptually relevant speech tracking in auditory and motor cortex reflects distinct linguistic features

During online speech processing, our brain tracks the acoustic fluctuations in speech at different timescales. Previous research has focused on generic timescales (for example, delta or theta bands) that are assumed to map onto linguistic features such as prosody or syllables. However, given the high intersubject variability in speaking patterns, such a generic association between the timescales of brain activity and speech properties can be ambiguous. Here, we analyse speech tracking in source-localised magnetoencephalographic data by directly focusing on timescales extracted from statistical regularities in our speech material. This revealed widespread significant tracking at the timescales of phrases (0.6–1.3 Hz), words (1.8–3 Hz), syllables (2.8–4.8 Hz), and phonemes (8–12.4 Hz). Importantly, when examining its perceptual relevance, we found stronger tracking for correctly comprehended trials in the left premotor (PM) cortex at the phrasal scale as well as in left middle temporal cortex at the word scale. Control analyses using generic bands confirmed that these effects were specific to the speech regularities in our stimuli. Furthermore, we found that the phase at the phrasal timescale coupled to power at beta frequency (13–30 Hz) in motor areas. This cross-frequency coupling presumably reflects top-down temporal prediction in ongoing speech perception. Together, our results reveal specific functional and perceptually relevant roles of distinct tracking and cross-frequency processes along the auditory–motor pathway

Alpha-band rhythms in visual task performance: phase-locking by rhythmic sensory stimulation

Oscillations are an important aspect of neuronal activity. Interestingly, oscillatory patterns are also observed in behaviour, such as in visual performance measures after the presentation of a brief sensory event in the visual or another modality. These oscillations in visual performance cycle at the typical frequencies of brain rhythms, suggesting that perception may be closely linked to brain oscillations. We here investigated this link for a prominent rhythm of the visual system (the alpha-rhythm, 8-12 Hz) by applying rhythmic visual stimulation at alpha-frequency (10.6 Hz), known to lead to a resonance response in visual areas, and testing its effects on subsequent visual target discrimination. Our data show that rhythmic visual stimulation at 10.6 Hz: 1) has specific behavioral consequences, relative to stimulation at control frequencies (3.9 Hz, 7.1 Hz, 14.2 Hz), and 2) leads to alpha-band oscillations in visual performance measures, that 3) correlate in precise frequency across individuals with resting alpha-rhythms recorded over parieto-occipital areas. The most parsimonious explanation for these three findings is entrainment (phase-locking) of ongoing perceptually relevant alpha-band brain oscillations by rhythmic sensory events. These findings are in line with occipital alpha-oscillations underlying periodicity in visual performance, and suggest that rhythmic stimulation at frequencies of intrinsic brain-rhythms can be used to reveal influences of these rhythms on task performance to study their functional roles

Being first matters: topographical representational similarity analysis of ERP signals reveals separate networks for audiovisual temporal binding depending on the leading sense

In multisensory integration, processing in one sensory modality is enhanced by complementary information from other modalities. Inter-sensory timing is crucial in this process as only inputs reaching the brain within a restricted temporal window are perceptually bound. Previous research in the audiovisual field has investigated various features of the temporal binding window (TBW), revealing asymmetries in its size and plasticity depending on the leading input (auditory-visual, AV; visual-auditory, VA). We here tested whether separate neuronal mechanisms underlie this AV-VA dichotomy in humans. We recorded high-density EEG while participants performed an audiovisual simultaneity judgment task including various AV/VA asynchronies and unisensory control conditions (visual-only, auditory-only) and tested whether AV and VA processing generate different patterns of brain activity. After isolating the multisensory components of AV/VA event-related potentials (ERPs) from the sum of their unisensory constituents, we run a time-resolved topographical representational similarity analysis (tRSA) comparing AV and VA ERP maps. Spatial cross-correlation matrices were built from real data to index the similarity between AV- and VA-maps at each time point (500ms window post-stimulus) and then correlated with two alternative similarity model matrices: AVmaps=VAmaps vs. AVmaps≠VAmaps. The tRSA results favored the AVmaps≠VAmaps model across all time points, suggesting that audiovisual temporal binding (indexed by synchrony perception) engages different neural pathways depending on the leading sense. The existence of such dual route supports recent theoretical accounts proposing that multiple binding mechanisms are implemented in the brain to accommodate different information parsing strategies in auditory and visual sensory systems