Group "Language and Computation in Neural Systems" and by the Netherlands Organization for Scientific Research (grant 016.Vidi.188.029).

It is undisputed that presenting a rhythmic stimulus leads to a measurable brain response that follows the rhythmic structure of this stimulus. What is still debated, however, is the question whether this brain response exclusively reflects a regular repetition of evoked responses, or whether it also includes entrained oscillatory activity. Here we systematically present evidence in favor of an involvement of entrained neural oscillations in the processing of rhythmic input while critically pointing out which questions still need to be addressed before this evidence could be considered conclusive. In this context, we also explicitly discuss the potential functional role of such entrained oscillations, suggesting that these stimulus-aligned oscillations reflect, and serve as, predictive processes, an idea often only implicitly assumed in the literature.

The role of oscillatory phase for perceptual and cognitive processes is being increasingly acknowledged. To date, little is known about the direct role of phase in categorical perception. Here we show in two separate experiments that the identification of ambiguous syllables that can either be perceived as /da/ or /ga/ is biased by the underlying oscillatory phase as measured with EEG and sensory entrainment to rhythmic stimuli. The measured phase difference in which perception is biased toward /da/ or /ga/ exactly matched the different temporal onset delays in natural audiovisual speech between mouth movements and speech sounds, which last 80 ms longer for /ga/ than for /da/. These results indicate the functional relationship between prestimulus phase and syllable identification, and signify that the origin of this phase relationship could lie in exposure and subsequent learning of unique audiovisual temporal onset differences.oscillations | phase | audiovisual | speech | temporal processing I n spoken language, visual mouth movements naturally precede the production of any speech sound, and therefore serve as a temporal prediction and detection cue for identifying spoken language (1) (but also see ref.2). Different syllables are characterized by unique visual-to-auditory temporal asynchronies (3, 4). For example, /ga/ has an 80-ms longer delay than /da/, and this difference aids categorical perception of these syllables (4). We propose that neuronal oscillations might carry the information to dissociate these syllables based on temporal differences. Multiple authors have proposed (5-7)-and it has been demonstrated empirically (7-9)-that at the onset of visual mouth movements, ongoing oscillations in auditory cortex align (see refs. 10-12 for nonspeech phase reset), providing a temporal reference frame for the auditory processing of subsequent speech sounds. Consequently, auditory signals fall on different phases of the aligned oscillation depending on the unique visualto-auditory temporal asynchrony, resulting in a consistent relationship between syllable identity and oscillatory phase.We hypothesized that this consistent "phase-syllable" relationship results in ongoing oscillatory phase biasing syllable perception. More specifically, the phase at which syllable perception is mostly biased should be proportional to the visual-toauditory temporal asynchrony found in natural speech. A naturally occurring /ga/ has an 80-ms longer visual-to-auditory onset difference than a naturally occurring /da/ (4). Consequently, the phase difference between perception bias toward /da/ and /ga/ should match 80 ms, which can only be established with an oscillation with a period greater than 80 ms, that is, any oscillation under 12.5 Hz. The apparent relevant oscillation range is therefore theta, with periods ranging between 111 and 250 ms (4-9 Hz). This oscillation range has already been proposed as a candidate to encode information, and seems specifically important for speech perception (13,14).To test this hypothesis of oscill...

No abstract

BackgroundVoluntary shifts of visuospatial attention are associated with a lateralization of parieto-occipital alpha power (7-13Hz), i.e. higher power in the hemisphere ipsilateral and lower power contralateral to the locus of attention. Recent noninvasive neuromodulation studies demonstrated that alpha power can be experimentally increased using transcranial alternating current stimulation (tACS).Objective/HypothesisWe hypothesized that tACS at alpha frequency over the left parietal cortex induces shifts of attention to the left hemifield. However, spatial attention shifts not only occur voluntarily (endogenous/ top-down), but also stimulus-driven (exogenous/ bottom-up). To study the task-specificity of the potential effects of tACS on attentional processes, we administered three conceptually different spatial attention tasks.Methods36 healthy volunteers were recruited from an academic environment. In two separate sessions, we applied either high-density tACS at 10Hz, or sham tACS, for 35–40 minutes to their left parietal cortex. We systematically compared performance on endogenous attention, exogenous attention, and stimulus detection tasks.ResultsIn the endogenous attention task, a greater leftward bias in reaction times was induced during left parietal 10Hz tACS as compared to sham. There were no stimulation effects in either the exogenous attention or the stimulus detection task.ConclusionThe study demonstrates that high-density tACS at 10Hz can be used to modulate visuospatial attention performance. The tACS effect is task-specific, indicating that not all forms of attention are equally susceptible to the stimulation.

Crossmodal binding usually relies on bottom-up stimulus characteristics such as spatial and temporal correspondence. However, in case of ambiguity the brain has to decide whether to combine or segregate sensory inputs. We hypothesise that widespread, subtle forms of synesthesia provide crossmodal mapping patterns which underlie and influence multisensory perception. Our aim was to investigate if such a mechanism plays a role in the case of pitch-size stimulus combinations. Using a combination of psychophysics and ERPs, we could show that despite violations of spatial correspondence, the brain specifically integrates certain stimulus combinations which are congruent with respect to our hypothesis of pitch-size synesthesia, thereby impairing performance on an auditory spatial localisation task (Ventriloquist effect). Subsequently, we perturbed this process by functionally disrupting a brain area known for its role in multisensory processes, the right intraparietal sulcus, and observed how the Ventriloquist effect was abolished, thereby increasing behavioural performance. Correlating behavioural, TMS and ERP results, we could retrace the origin of the synesthestic pitch-size mappings to a right intraparietal involvement around 250 ms. The results of this combined psychophysics, TMS and ERP study provide evidence for shifting the current viewpoint on synesthesia more towards synesthesia being at the extremity of a spectrum of normal, adaptive perceptual processes, entailing close interplay between the different sensory systems. Our results support this spectrum view of synesthesia by demonstrating that its neural basis crucially depends on normal multisensory processes.

Temporal structure in the environment often has predictive value for anticipating the occurrence of forthcoming events. In this study we investigated the influence of two types of predictive temporal information on the perception of near-threshold auditory stimuli: 1) intrinsic temporal rhythmicity within an auditory stimulus stream and 2) temporally-predictive visual cues. We hypothesized that combining predictive temporal information within- and across-modality should decrease the threshold at which sounds are detected, beyond the advantage provided by each information source alone. Two experiments were conducted in which participants had to detect tones in noise. Tones were presented in either rhythmic or random sequences and were preceded by a temporally predictive visual signal in half of the trials. We show that detection intensities are lower for rhythmic (vs. random) and audiovisual (vs. auditory-only) presentation, independent from response bias, and that this effect is even greater for rhythmic audiovisual presentation. These results suggest that both types of temporal information are used to optimally process sounds that occur at expected points in time (resulting in enhanced detection), and that multiple temporal cues are combined to improve temporal estimates. Our findings underscore the flexibility and proactivity of the perceptual system which uses within- and across-modality temporal cues to anticipate upcoming events and process them optimally.

Many environmental stimuli contain temporal regularities, a feature that can help predict forthcoming input. Phase locking (entrainment) of ongoing low-frequency neuronal oscillations to rhythmic stimuli is proposed as a potential mechanism for enhancing neuronal responses and perceptual sensitivity, by aligning high-excitability phases to events within a stimulus stream. Previous experiments show that rhythmic structure has a behavioral benefit even when the rhythm itself is below perceptual detection thresholds (ten Oever et al., 2014). It is not known whether this "inaudible" rhythmic sound stream also induces entrainment. Here we tested this hypothesis using magnetoencephalography and electrocorticography in humans to record changes in neuronal activity as subthreshold rhythmic stimuli gradually became audible. We found that significant phase locking to the rhythmic sounds preceded participants' detection of them. Moreover, no significant auditory-evoked responses accompanied this prethreshold entrainment. These auditory-evoked responses, distinguished by robust, broad-band increases in intertrial coherence, only appeared after sounds were reported as audible. Taken together with the reduced perceptual thresholds observed for rhythmic sequences, these findings support the proposition that entrainment of low-frequency oscillations serves a mechanistic role in enhancing perceptual sensitivity for temporally predictive sounds. This framework has broad implications for understanding the neural mechanisms involved in generating temporal predictions and their relevance for perception, attention, and awareness. The environment is full of rhythmically structured signals that the nervous system can exploit for information processing. Thus, it is important to understand how the brain processes such temporally structured, regular features of external stimuli. Here we report the alignment of slowly fluctuating oscillatory brain activity to external rhythmic structure before its behavioral detection. These results indicate that phase alignment is a general mechanism of the brain to process rhythmic structure and can occur without the perceptual detection of this temporal structure.