Keiichi Kitajo

Synchronization of spontaneous eyeblinks while viewing video stories

Blinks are generally suppressed during a task that requires visual attention and tend to occur immediately before or after the task when the timing of its onset and offset are explicitly given. During the viewing of video stories, blinks are expected to occur at explicit breaks such as scene changes. However, given that the scene length is unpredictable, there should also be appropriate timing for blinking within a scene to prevent temporal loss of critical visual information. Here, we show that spontaneous blinks were highly synchronized between and within subjects when they viewed the same short video stories, but were not explicitly tied to the scene breaks. Synchronized blinks occurred during scenes that required less attention such as at the conclusion of an action, during the absence of the main character, during a long shot and during repeated presentations of a similar scene. In contrast, blink synchronization was not observed when subjects viewed a background video or when they listened to a story read aloud. The results suggest that humans share a mechanism for controlling the timing of blinks that searches for an implicit timing that is appropriate to minimize the chance of losing critical information while viewing a stream of visual events.

Dynamic links between theta executive functions and alpha storage buffers in auditory and visual working memory

Working memory (WM) tasks require not only distinct functions such as a storage buffer and central executive functions, but also coordination among these functions. Neuroimaging studies have revealed the contributions of different brain regions to different functional roles in WM tasks; however, little is known about the neural mechanism governing their coordination. Electroencephalographic (EEG) rhythms, especially theta and alpha, are known to appear over distributed brain regions during WM tasks, but the rhythms associated with task‐relevant regional coupling have not been obtained thus far. In this study, we conducted time–frequency analyses for EEG data in WM tasks that include manipulation periods and memory storage buffer periods. We used both auditory WM tasks and visual WM tasks. The results successfully demonstrated function‐specific EEG activities. The frontal theta amplitudes increased during the manipulation periods of both tasks. The alpha amplitudes increased during not only the manipulation but also the maintenance periods in the temporal area for the auditory WM and the parietal area for the visual WM. The phase synchronization analyses indicated that, under the relevant task conditions, the temporal and parietal regions show enhanced phase synchronization in the theta bands with the frontal region, whereas phase synchronization between theta and alpha is significantly enhanced only within the individual areas. Our results suggest that WM task‐relevant brain regions are coordinated by distant theta synchronization for central executive functions, by local alpha synchronization for the memory storage buffer, and by theta–alpha coupling for inter‐functional integration.

Noise-induced large-scale phase synchronization of human-brain activity associated with behavioural stochastic resonance

We demonstrate that both detection of weak visual signals to the right eye and phase synchronization of electro-encephalogram (EEG) signals from widely separated areas of the human brain are increased by addition of weak visual noise to the left eye. We found a close relationship between the resulting noise-induced changes in behavioural performance and the similarly resulting changes in phase synchronization between widely separated brain areas. These results imply that noise-induced large-scale neural synchronization may play a significant role in information transmission in the brain.

Internal noise determines external stochastic resonance in visual perception

We provide the first experimental evidence that the internal noise level determines whether external noise can enhance the detectability of a weak signal. We conduct a visual detection experiment in the absence and presence of visual noise. We define three indices of external stochastic resonance effects, consider the spread of the psychometric function without external noise as an internal noise level index, and find that the indices of external stochastic resonance effects negatively correlate with the internal noise level index. Our results suggest that external stochastic resonance depends not only on the external but also on the internal noise level.

Transcranial magnetic stimulation-induced global propagation of transient phase resetting associated with directional information flow

Electroencephalogram (EEG) phase synchronization analyses can reveal large-scale communication between distant brain areas. However, it is not possible to identify the directional information flow between distant areas using conventional phase synchronization analyses. In the present study, we applied transcranial magnetic stimulation (TMS) to the occipital area in subjects who were resting with their eyes closed, and analyzed the spatial propagation of transient TMS-induced phase resetting by using the transfer entropy (TE), to quantify the causal and directional flow of information. The time-frequency EEG analysis indicated that the theta (5 Hz) phase locking factor (PLF) reached its highest value at the distant area (the motor area in this study), with a time lag that followed the peak of the transient PLF enhancements of the TMS-targeted area at the TMS onset. Phase-preservation index (PPI) analyses demonstrated significant phase resetting at the TMS-targeted area and distant area. Moreover, the TE from the TMS-targeted area to the distant area increased clearly during the delay that followed TMS onset. Interestingly, the time lags were almost coincident between the PLF and TE results (152 vs. 165 ms), which provides strong evidence that the emergence of the delayed PLF reflects the causal information flow. Such tendencies were observed only in the higher-intensity TMS condition, and not in the lower-intensity or sham TMS conditions. Thus, TMS may manipulate large-scale causal relationships between brain areas in an intensity-dependent manner. We demonstrated that single-pulse TMS modulated global phase dynamics and directional information flow among synchronized brain networks. Therefore, our results suggest that single-pulse TMS can manipulate both incoming and outgoing information in the TMS-targeted area associated with functional changes.

Fronto-parietal and fronto-temporal theta phase synchronization for visual and auditory-verbal working memory

In humans, theta phase (4–8 Hz) synchronization observed on electroencephalography (EEG) plays an important role in the manipulation of mental representations during working memory (WM) tasks; fronto-temporal synchronization is involved in auditory-verbal WM tasks and fronto-parietal synchronization is involved in visual WM tasks. However, whether or not theta phase synchronization is able to select the to-be-manipulated modalities is uncertain. To address the issue, we recorded EEG data from subjects who were performing auditory-verbal and visual WM tasks; we compared the theta synchronizations when subjects performed either auditory-verbal or visual manipulations in separate WM tasks, or performed both two manipulations in the same WM task. The auditory-verbal WM task required subjects to calculate numbers presented by an auditory-verbal stimulus, whereas the visual WM task required subjects to move a spatial location in a mental representation in response to a visual stimulus. The dual WM task required subjects to manipulate auditory-verbal, visual, or both auditory-verbal and visual representations while maintaining auditory-verbal and visual representations. Our time-frequency EEG analyses revealed significant fronto-temporal theta phase synchronization during auditory-verbal manipulation in both auditory-verbal and auditory-verbal/visual WM tasks, but not during visual manipulation tasks. Similarly, we observed significant fronto-parietal theta phase synchronization during visual manipulation tasks, but not during auditory-verbal manipulation tasks. Moreover, we observed significant synchronization in both the fronto-temporal and fronto-parietal theta signals during simultaneous auditory-verbal/visual manipulations. These findings suggest that theta synchronization seems to flexibly connect the brain areas that manipulate WM.

Neural dynamics in motor preparation: From phase-mediated global computation to amplitude-mediated local computation

Oscillatory activity plays a critical role in the brain. Here, we illustrate the dynamics of neural oscillations in the motor system of the brain. We used a non-directional cue to instruct participants to prepare a motor response with either the left or the right hand and recorded electroencephalography during the preparation of the response. Consistent with previous findings, the amplitude of alpha-band (8-14Hz) oscillations significantly decreased over the motor region contralateral to the hand prepared for the response. Prior to this decrease, there were a number of inter-regional phase synchronies at lower frequencies (2-4Hz; delta band). Cross-frequency coupling was quantified to further explore the direct link between alpha amplitudes and delta synchrony. The cross-frequency coupling of showed response-specific modulation, whereby the motor region contralateral to the preparation hand exhibited an increase in coupling relative to the baseline. The amplitude of alpha oscillations had an unpreferred and a preferred delta phase, in which the amplitude was modulated negatively and positively, respectively. Given the amplitude of alpha-band oscillations decreased over the analyzed period, the alpha amplitude might be down-regulated by the phase-amplitude coupling, although we do not have direct evidence for that. Taken together, these results show global-to-local computation in the motor system, which started from inter-regional delta phase synchrony and ended at an effector-specific decrease in the amplitude of alpha-band oscillations, with phase-amplitude coupling connecting both computations.

Transient process of cortical activity during Necker cube perception: from local clusters to global synchrony

Background It has been discussed that neural phase-synchrony across distant cortical areas (or global phase-synchrony) was correlated with various aspects of consciousness. The generating process of the synchrony, however, remains largely unknown. As a first step, we investigate transient process of global phase-synchrony, focusing on phase-synchronized clusters. We hypothesize that the phase-synchronized clusters are dynamically organized before global synchrony and clustering patterns depend on perceptual conditions. Methods In an EEG study, Kitajo reported that phase-synchrony across distant cortical areas was selectively enhanced by top-down attention around 4 Hz in Necker cube perception. Here, we further analyzed the phase-synchronized clusters using hierarchical clustering which sequentially binds up the nearest electrodes based on similarity of phase locking between the cortical signals. First, we classified dominant components of the phase-synchronized clusters over time. We then investigated how the phase-synchronized clusters change with time, focusing on their size and spatial structure. Results Phase-locked clusters organized a stable spatial pattern common to the perceptual conditions. In addition, the phase-locked clusters were modulated transiently depending on the perceptual conditions and the time from the perceptual switch. When top-down attention succeeded in switching perception as subjects intended, independent clusters at frontal and occipital areas grew to connect with each other around the time of the perceptual switch. However, the clusters in the occipital and left parietal areas remained divided when top-down attention failed in switching perception. When no primary biases exist, the cluster in the occipital area grew to its maximum at the time of the perceptual switch within the occipital area. Conclusions Our study confirmed the existence of stable phase-synchronized clusters. Furthermore, these clusters were transiently connected with each other. The connecting pattern depended on subjects’ internal states. These results suggest that subjects’ attentional states are associated with distinct spatio-temporal patterns of the phase-locked clusters.

Detection of phase synchronization in EEG with Bivariate Empirical Mode Decomposition

In recent years, the phase synchronization phenomenon in the electroencephalograph (EEG) has been widely used to observe interactions between separate areas of the cortex. However, the traditional coherence to measure the phase synchronization need target signals to be stationary. In this paper, we propose a technique to measure the phase synchrony of non-stationary signals by the Phase Locking Value (PLV) with Hilbert transform and the Bivariate Empirical Mode Decomposition (BEMD). We analyzed the phase synchronization of EEG signals which were recorded during Dynamical Dot Quartet (DDQ) tasks using the conventional method and the proposed method. The analysis result suggests that proposed method more suitable for detecting the phase synchrony during the DDQ tasks than the conventional methods.

TMS-induced theta phase synchrony reveals a bottom-up network in working memory

Global theta phase synchronization between the frontal and sensory areas has been suggested to connect the relevant areas for executive processes of working memory (WM). However, little is known regarding network directionality (i.e. top-down or bottom-up) of this interaction. To address the issue, the present study conducted transcranial magnetic stimulation (TMS)-electroencephalography (EEG) experiment during WM tasks. Results showed that TMS-induced increases in theta phase synchronization were observed only when TMS was delivered to the sensory areas but not the frontal area. These findings suggest that network directionality represented in WM is bottom-up rather than top-down.