Frequency-Dependent Cortical Interactions during Semantic Processing: An Electrocorticogram Cross-spectrum Analysis Using a Semantic Space Model.

Convergent evidence has demonstrated that semantics are represented by the interaction between a multimodal semantic hub at the anterior temporal lobe (ATL) and other modality-specific association cortical areas. Electrocorticogram (ECoG) recording with high spatiotemporal resolutions is efficient in evaluating such cortical interactions; however, this has not been a focus of preceding studies. The present study evaluated cortical interactions during picture naming using a novel ECoG cross-spectrum analysis, which was formulated from a computational simulation of neuronal networks and combined with a vector space model of semantics. The results clarified three types of frequency-dependent cortical networks: 1) an earlier-period (0.2-0.8 s from stimulus onset) high-gamma-band (90-150 Hz) network with a hub at the posterior fusiform gyrus, 2) a later-period (0.4-1.0 s) beta-band (15-40 Hz) network with multiple hubs at the ventral ATL and posterior middle temporal gyrus, and 3) a pre-articulation theta-band (4-7 Hz) network distributed over widely located cortical regions. These results suggest that frequency-dependent cortical interactions can characterize the underlying processes of semantic cognition, and the beta-band network with a hub at the ventral ATL is especially associated with the formation of semantic representation.

A dynamical systems approach for estimating phase interactions between rhythms of different frequencies from experimental data.

Synchronization of neural oscillations as a mechanism of brain function is attracting increasing attention. Neural oscillation is a rhythmic neural activity that can be easily observed by noninvasive electroencephalography (EEG). Neural oscillations show the same frequency and cross-frequency synchronization for various cognitive and perceptual functions. However, it is unclear how this neural synchronization is achieved by a dynamical system. If neural oscillations are weakly coupled oscillators, the dynamics of neural synchronization can be described theoretically using a phase oscillator model. We propose an estimation method to identify the phase oscillator model from real data of cross-frequency synchronized activities. The proposed method can estimate the coupling function governing the properties of synchronization. Furthermore, we examine the reliability of the proposed method using time-series data obtained from numerical simulation and an electronic circuit experiment, and show that our method can estimate the coupling function correctly. Finally, we estimate the coupling function between EEG oscillation and the speech sound envelope, and discuss the validity of these results.

Cortical networks dynamically emerge with the interplay of slow and fast oscillations for memory of a natural scene.

Neural oscillations are crucial for revealing dynamic cortical networks and for serving as a possible mechanism of inter-cortical communication, especially in association with mnemonic function. The interplay of the slow and fast oscillations might dynamically coordinate the mnemonic cortical circuits to rehearse stored items during working memory retention. We recorded simultaneous EEG-fMRI during a working memory task involving a natural scene to verify whether the cortical networks emerge with the neural oscillations for memory of the natural scene. The slow EEG power was enhanced in association with the better accuracy of working memory retention, and accompanied cortical activities in the mnemonic circuits for the natural scene. Fast oscillation showed a phase-amplitude coupling to the slow oscillation, and its power was tightly coupled with the cortical activities for representing the visual images of natural scenes. The mnemonic cortical circuit with the slow neural oscillations would rehearse the distributed natural scene representations with the fast oscillation for working memory retention. The coincidence of the natural scene representations could be obtained by the slow oscillation phase to create a coherent whole of the natural scene in the working memory.

Dynamic parieto-premotor network for mental image transformation revealed by simultaneous EEG and fMRI measurement.

Previous studies have suggested that the posterior parietal cortices and premotor areas are involved in mental image transformation. However, it remains unknown whether these regions really cooperate to realize mental image transformation. In this study, simultaneous EEG and fMRI were performed to clarify the spatio-temporal properties of neural networks engaged in mental image transformation. We adopted a modified version of the mental clock task used by Sack et al. [Sack, A. T., Camprodon, J. A., Pascual-Leone, A., & Goebel, R. The dynamics of interhemispheric compensatory processes in mental imagery. Science, 308, 702-704, 2005; Sack, A. T., Sperling, J. M., Prvulovic, D., Formisano, E., Goebel, R., Di Salle, F., et al. Tracking the mind's image in the brain II: Transcranial magnetic stimulation reveals parietal asymmetry in visuospatial imagery. Neuron, 35, 195-204, 2002]. In the modified mental clock task, participants mentally rotated clock hands from the position initially presented at a learned speed for various durations. Subsequently, they matched the position to the visually presented clock hands. During mental rotation of the clock hands, we observed significant beta EEG suppression with respect to the amount of mental rotation at the right parietal electrode. The beta EEG suppression accompanied activity in the bilateral parietal cortices and left premotor cortex, representing a dynamic cortical network for mental image transformation. These results suggest that motor signals from the premotor area were utilized for mental image transformation in the parietal areas and for updating the imagined clock hands represented in the right posterior parietal cortex.

Neural activity related to productive vocabulary knowledge effects during second language comprehension.

Second language learners and educators often believe that improving one's listening ability hinges on acquiring an extensive vocabulary and engaging in thorough listening practice. Our previous study suggested that listening comprehension is also impacted by the ability to produce vocabulary. Nevertheless, it remained uncertain whether quick comprehension could be attributed to a simple acceleration of processing or to changes in neural activity. To identify neural activity changes during sentence listening comprehension according to different levels of lexical knowledge (productive, only comprehensive, uncomprehensive), we measured participants' electrical activity in the brain via electroencephalography (EEG) and conducted a time-frequency-based EEG power analysis. Additionally, we employed a decoding model to verify the predictability of vocabulary knowledge levels based on neural activity. The decoding results showed that EEG activity could discriminate between listening to sentences containing phrases that include productive knowledge and ones without. The positive impact of productive vocabulary knowledge on sentence comprehension, driven by distinctive neural processing during sentence comprehension, was unequivocally evident. Our study emphasizes the importance of productive vocabulary knowledge acquisition to enhance the process of second language listening comprehension.

The effect of productive vocabulary knowledge on second language comprehension.

Second language learners tend to focus more on learning the meaning of vocabulary than on how to use it in their speech and writing. Although comprehensive vocabulary knowledge is necessary for understanding sentences, productive vocabulary knowledge may also have a positive impact on sentence comprehension. Most studies examining the relationship between production and comprehension have compared these abilities between participants or evaluated unrelated criteria between tasks, which may be insufficient for examining the direct effects of productive knowledge on sentence comprehension. Our study investigates changes in sentence comprehension speed during listening, and we used a within-subjects comparison to examine the effect of productive vocabulary knowledge or the lack thereof. We applied generalized linear mixed models to investigate productive vocabulary knowledge effects by partialing out other influential factors, such as confidence, frequency of target words, stimulus duration, and individual differences. The results showed that the sentences with a producible phrase were processed faster than the sentences that included phrases with only comprehensive knowledge or no comprehension. The effect of productive vocabulary knowledge on the speed of sentence comprehension was directly examined with a within-subject comparison, and its contribution was clearly found. This study emphasizes the value of productive vocabulary knowledge acquisition in enhancing the speed of sentence comprehension.

Cortical dynamics of human scalp EEG origins in a visually guided motor execution.

The EEG mu rhythm is often used as an index of activation in the sensorimotor cortex. However, the blur caused by volume conduction makes it difficult to identify the exact origin of the EEG rhythm in the brain using only the human scalp EEG. In this study, simultaneous fMRI and EEG measurements were performed during a visually guided motor execution task in order to investigate whether the mu rhythm in the scalp EEG is an indication of the activity in the sensorimotor cortex. In addition, a new method was proposed for reconstruction of the cortical EEG activity through the fusion of fMRI and EEG data. A suppression of mu rhythm appeared around the lateral central electrode sites, just above the sensorimotor cortex, in association with the activity in that region. During a visually guided motor execution task, the alpha rhythms at the occipital electrode sites and the alpha rhythm at the central electrode sites also showed a correlation with the fMRI signal in the occipital and the supplementary motor cortices, respectively. This method allows the investigation of the scalp EEG origin with the spatial precision of fMRI, while retaining dynamic properties of the cortex with the temporal precision of EEG.

Neuronal ensemble for visual working memory via interplay of slow and fast oscillations.

The current focus of studies on neural entities for memory maintenance is on the interplay between fast neuronal oscillations in the gamma band and slow oscillations in the theta or delta band. The hierarchical coupling of slow and fast oscillations is crucial for the rehearsal of sensory inputs for short-term storage, as well as for binding sensory inputs that are represented in spatially segregated cortical areas. However, no experimental evidence for the binding of spatially segregated information has yet been presented for memory maintenance in humans. In the present study, we actively manipulated memory maintenance performance with an attentional blink procedure during human scalp electroencephalography (EEG) recordings and identified that slow oscillations are enhanced when memory maintenance is successful. These slow oscillations accompanied fast oscillations in the gamma frequency range that appeared at spatially segregated scalp sites. The amplitude of the gamma oscillation at these scalp sites was simultaneously enhanced at an EEG phase of the slow oscillation. Successful memory maintenance appears to be achieved by a rehearsal of sensory inputs together with a coordination of distributed fast oscillations at a preferred timing of the slow oscillations.

The Role of Temporal Contingency and Integrity of Visual Inputs in the Sense of Agency: A Psychophysical Study.

The sense of agency is a subjective feeling that one's own actions drive action outcomes. Previous studies have focused primarily on the temporal contingency between actions and sensory inputs as a possible mechanism for the sense of agency. However, the contribution of the integrity of visual inputs has not been systematically addressed. In the current study, we developed a psychophysical task to examine the role of visual inputs as well as temporal contingencies toward the sense of agency. Specifically, participants were required to track a target on a sinusoidal curve on a computer screen. Visual integrity of sensory inputs was manipulated by gradually occluding a computer cursor, and participants were asked to report the sense of agency on a nine-point Likert scale. Temporal contingency was manipulated by varying the delay between finger movements on a touchpad and cursor movements. The results showed that the sense of agency was influenced by both visual integrity and temporal contingency. These results are discussed in the context of current models that have proposed that the sense of agency emerges from the comparison of visual inputs with motor commands.

Nonimmersive virtual reality mirror visual feedback therapy and its application for the treatment of complex regional pain syndrome: an open-label pilot study.

OBJECTIVE: Chronic pain conditions such as phantom limb pain and complex regional pain syndrome are difficult to treat, and traditional pharmacological treatment and invasive neural block are not always effective. Plasticity in the central nervous system occurs in these conditions and may be associated with pain. Mirror visual feedback therapy aims to restore normal cortical organization and is applied in the treatment of chronic pain conditions. However, not all patients benefit from this treatment. Virtual reality technology is increasingly attracting attention for medical application, including as an analgesic modality. An advanced mirror visual feedback system with virtual reality technology may have increased analgesic efficacy and benefit a wider patient population. In this preliminary work, we developed a virtual reality mirror visual feedback system and applied it to the treatment of complex regional pain syndrome. DESIGN: A small open-label case series. Five patients with complex regional pain syndrome received virtual reality mirror visual feedback therapy once a week for five to eight sessions on an outpatient basis. Patients were monitored for continued medication use and pain intensity. RESULTS: Four of the five patients showed >50% reduction in pain intensity. Two of these patients ended their visits to our pain clinic after five sessions. CONCLUSION: Our results indicate that virtual reality mirror visual feedback therapy is a promising alternative treatment for complex regional pain syndrome. Further studies are necessary before concluding that analgesia provided from virtual reality mirror visual feedback therapy is the result of reversing maladaptive changes in pain perception.

Successful Encoding during Natural Reading Is Associated with Fixation-Related Potentials and Large-Scale Network Deactivation.

Reading literature (e.g., an entire book) is an enriching experience that qualitatively differs from reading a single sentence; however, the brain dynamics of such context-dependent memory remains unclear. This study aimed to elucidate mnemonic neural dynamics during natural reading of literature by performing electroencephalogram (EEG) and functional magnetic resonance imaging (fMRI). Brain activities of human participants recruited on campus were correlated with their subsequent memory, which was quantified by semantic correlation between the read text and reports subsequently written by them based on state of the art natural language processing procedures. The results of the EEG data analysis showed a significant positive relationship between subsequent memory and fixation-related EEG. Sentence-length and paragraph-length mnemonic processes were associated with N1-P2 and P3 fixation-related potential (FRP) components and fixation-related θ-band (4-8 Hz) EEG power, respectively. In contrast, the results of fMRI analysis showed a significant negative relationship between subsequent memory and blood oxygenation level-dependent (BOLD) activation. Sentence-length and paragraph-length mnemonic processes were associated with networks of regions forming part of the salience network and the default mode network (DMN), respectively. Taken together with the EEG results, these memory-related deactivations in the salience network and the DMN were thought to reflect the reading of sentences characterized by low mnemonic load and the suppression of task-irreverent thoughts, respectively. It was suggested that the context-dependent mnemonic process during literature reading requires large-scale network deactivation, which might reflect coordination of a range of voluntary processes during reading.

Open monitoring meditation reduces the involvement of brain regions related to memory function.

Mindfulness meditation consists of focused attention meditation (FAM) and open monitoring meditation (OMM), both of which reduce activation of the default mode network (DMN) and mind-wandering. Although it is known that FAM requires intentional focused attention, the mechanisms of OMM remain largely unknown. To investigate this, we examined striatal functional connectivity in 17 experienced meditators (mean total practice hours = 920.6) during pre-resting, meditation, and post-resting states comparing OMM with FAM, using functional magnetic resonance imaging. Both FAM and OMM reduced functional connectivity between the striatum and posterior cingulate cortex, which is a core hub region of the DMN. Furthermore, OMM reduced functional connectivity of the ventral striatum with both the visual cortex related to intentional focused attention in the attentional network and retrosplenial cortex related to memory function in the DMN. In contrast, FAM increased functional connectivity in these regions. Our findings suggest that OMM reduces intentional focused attention and increases detachment from autobiographical memory. This detachment may play an important role in non-judgmental and non-reactive attitude during OMM. These findings provide new insights into the mechanisms underlying the contribution of OMM to well-being and happiness.

Ongoing slow oscillatory phase modulates speech intelligibility in cooperation with motor cortical activity.

Neural oscillation is attracting attention as an underlying mechanism for speech recognition. Speech intelligibility is enhanced by the synchronization of speech rhythms and slow neural oscillation, which is typically observed as human scalp electroencephalography (EEG). In addition to the effect of neural oscillation, it has been proposed that speech recognition is enhanced by the identification of a speaker's motor signals, which are used for speech production. To verify the relationship between the effect of neural oscillation and motor cortical activity, we measured scalp EEG, and simultaneous EEG and functional magnetic resonance imaging (fMRI) during a speech recognition task in which participants were required to recognize spoken words embedded in noise sound. We proposed an index to quantitatively evaluate the EEG phase effect on behavioral performance. The results showed that the delta and theta EEG phase before speech inputs modulated the participant's response time when conducting speech recognition tasks. The simultaneous EEG-fMRI experiment showed that slow EEG activity was correlated with motor cortical activity. These results suggested that the effect of the slow oscillatory phase was associated with the activity of the motor cortex during speech recognition.

Top-down and bottom-up attention cause the ventriloquism effect with distinct electroencephalography modulations.

The ventriloquism effect is a critical phenomenon for understanding the underlying mechanisms of multisensory integration. Cross-modal spatial attention causes a distortion of sound localization, although the neural basis of the effect remains an unanswered question. We hypothesized that top-down and bottom-up visual-spatial attention causes the ventriloquism effect with different modulations of ongoing neural oscillation. To test this hypothesis, human scalp electroencephalography (EEG) was measured during a sound localization task. Top-down attention suppressed the EEG amplitude in the alpha frequency (10 Hz) over the contralateral temporal electrode sites to visual cue hemifields. Bottom-up attention shifted the EEG phase to the theta frequency (7 Hz), rather than suppressing the amplitude. Two different neural mechanisms of ongoing neural oscillation contributed toward the ventriloquism effect, with different spatial attention.

Increased BOLD Signals Elicited by High Gamma Auditory Stimulation of the Left Auditory Cortex in Acute State Schizophrenia.

Recent MRI studies have shown that schizophrenia is characterized by reductions in brain gray matter, which progress in the acute state of the disease. Cortical circuitry abnormalities in gamma oscillations, such as deficits in the auditory steady state response (ASSR) to gamma frequency (>30-Hz) stimulation, have also been reported in schizophrenia patients. In the current study, we investigated neural responses during click stimulation by BOLD signals. We acquired BOLD responses elicited by click trains of 20, 30, 40 and 80-Hz frequencies from 15 patients with acute episode schizophrenia (AESZ), 14 symptom-severity-matched patients with non-acute episode schizophrenia (NASZ), and 24 healthy controls (HC), assessed via a standard general linear-model-based analysis. The AESZ group showed significantly increased ASSR-BOLD signals to 80-Hz stimuli in the left auditory cortex compared with the HC and NASZ groups. In addition, enhanced 80-Hz ASSR-BOLD signals were associated with more severe auditory hallucination experiences in AESZ participants. The present results indicate that neural over activation occurs during 80-Hz auditory stimulation of the left auditory cortex in individuals with acute state schizophrenia. Given the possible association between abnormal gamma activity and increased glutamate levels, our data may reflect glutamate toxicity in the auditory cortex in the acute state of schizophrenia, which might lead to progressive changes in the left transverse temporal gyrus.

A long-range cortical network emerging with theta oscillation in a mental task.

Human scalp EEG has demonstrated that global coherence among distant areas increases during cognitive tasks, suggesting that oscillating neural activities work to generate global neuronal assemblies for cognitive functions. The theta oscillation in a frequency range of 4-8 Hz with large amplitudes which emerges during mental tasks around the frontal midline region is called fm theta. If theta oscillation concerns the global neuronal assemblies, fm theta should be associated with regional activities that depend on task conditions. In the present study, we examine the EEG-related brain activities by developing simultaneous EEG and fMRI during a mental calculation task. EEG-related negative BOLD was dominant over anterior medial regions, suggesting a major contribution of negative BOLD to fm theta. Negative and positive BOLD were found over distant regions. Functional connectivity analyses revealed that the connectivity varied remarkably according to mental conditions. In the rest condition, the connectivity was localized, whereas in the task condition, a long-range coherent network was formed by the anterior midline, posterior cingulate and right middle temporal cortices with linking between the right middle temporal and left lateral cortices during numerical processing. Further EEG analyses indicate that the long-range coherent network executing cognitive functions is coordinated in the time window of theta oscillations.

Traveling EEG slow oscillation along the dorsal attention network initiates spontaneous perceptual switching.

An ambiguous figure such as the Necker cube causes spontaneous perceptual switching (SPS). The mechanism of SPS in multistable perception has not yet been determined. Although early psychological studies suggested that SPS may be caused by fatigue or satiation of orientation, the neural mechanism of SPS is still unknown. Functional magnetic resonance imaging (fMRI) has shown that the dorsal attention network (DAN), which mainly controls voluntary attention, is involved in bistable perception of the Necker cube. To determine whether neural dynamics along the DAN cause SPS, we performed simultaneous electroencephalography (EEG) and fMRI during an SPS task with the Necker cube, with every SPS reported by pressing a button. This EEG-fMRI integrated analysis showed that (a) 3-4 Hz spectral EEG power modulation at fronto-central, parietal, and centro-parietal electrode sites sequentially appeared from 750 to 350 ms prior to the button press; and (b) activations correlating with the EEG modulation traveled along the DAN from the frontal to the parietal regions. These findings suggest that slow oscillation initiates SPS through global dynamics along the attentional system such as the DAN.

Subsequent memory-dependent EEG theta correlates to parahippocampal blood oxygenation level-dependent response.

The 4-12 Hz (theta rhythm)-dependent neural dynamics play a fundamental role in the memory formation of the rat hippocampus. Although the power of human scalp electroencephalography theta (EEG theta) is known to be associated with a hippocampus-dependent memory encoding, it remains unclear whether the human hippocampus uses theta rhythm. In this study, we aim to identify the scalp EEG theta-related neural regions during memory encoding by using a simultaneous EEG-functional magnetic resonance imaging recording. We showed that the parahippocampal and the medial frontal and posterior regions were significantly correlated to subsequent memory-dependent EEG theta power. This evidence suggests that the human parahippocampal region and associated structures use theta rhythm during hippocampal memory encoding as in rodents.

Human cortical circuits for central executive function emerge by theta phase synchronization.

Dynamic networking of brain regions is suggested to be one of the key factors involved in various brain computations. Central executive function typically requires instantaneous coordination among the medial prefrontal regions and other distant regions, depending on the on-going task situation. In human scalp-recorded electroencephalography (EEG), the medial prefrontal area is estimated to be the current source of the theta rhythm, while there is no direct evidence that the theta rhythm is involved in the dynamic networking of central executive circuits. Here we hypothesize that the central executive circuit over the prefrontal and task-related cortices is dynamically linked by theta synchronization. By using simultaneous functional magnetic resonance imaging (fMRI) and EEG, we elucidated cortical circuits emerging with theta phase synchronization during free pacing repeated subtraction. Theta phase synchronization in the scalp EEG was found to emerge at two major clusters of electrode pairs, between the right frontal and left parietal sites and between the frontal and right parietal sites. The phase synchronization of two clusters is accompanied by fMRI responses in the cortical regions responsible for central executive function, working memory, visual imagery and cognitive action sequence. Here we report the first evidence that theta phase synchronization dynamically coordinates the central executive circuits, including the medial prefrontal cortex and relevant cortical regions.