Temporal dynamics of the knowledge-mediated visual disambiguation process in humans: a magnetoencephalography study.

Disambiguation of a noisy visual scene with prior knowledge is an indispensable task of the visual system. To adequately adapt to a dynamically changing visual environment full of noisy visual scenes, the implementation of knowledge-mediated disambiguation in the brain is imperative and essential for proceeding as fast as possible under the limited capacity of visual image processing. However, the temporal profile of the disambiguation process has not yet been fully elucidated in the brain. The present study attempted to determine how quickly knowledge-mediated disambiguation began to proceed along visual areas after the onset of a two-tone ambiguous image using magnetoencephalography with high temporal resolution. Using the predictive coding framework, we focused on activity reduction for the two-tone ambiguous image as an index of the implementation of disambiguation. Source analysis revealed that a significant activity reduction was observed in the lateral occipital area at approximately 120 ms after the onset of the ambiguous image, but not in preceding activity (about 115 ms) in the cuneus when participants perceptually disambiguated the ambiguous image with prior knowledge. These results suggested that knowledge-mediated disambiguation may be implemented as early as approximately 120 ms following an ambiguous visual scene, at least in the lateral occipital area, and provided an insight into the temporal profile of the disambiguation process of a noisy visual scene with prior knowledge.

Cortical theta phase synchronization involved in mismatch-driven perceptual alternation in binocular rivalry.

When two conflicting images are presented to each eye, a phenomenon called binocular rivalry occurs in which we initially perceive one image, and then our perception switches to the other over time. An enhancement of θ-band phase coherence in visual mismatch oscillatory response (vMOR) is reported to be involved in the facilitation of perceptual alternation when the deviant stimulus is presented unconsciously. In this study, we investigated the modulation effect of θ-band transcranial alternating current stimulation (tACS) on perceptual alternation in binocular rivalry, with a focus on its relationship with the θ-band vMOR. The results showed that tACS had no significant effect on the mean proportion of perceptual alternation. Analyzing the differential effects of the modulation, however, we found a positive correlation between the increase in inter-trial phase coherence of the vMOR and the promotion of perceptual alternation under the unconscious deviant condition. Additionally, our findings indicate that the θ-band phase synchrony between frontal and occipital electrode sides, as measured by the phase lag index, is implicated in perceptual alternation, with an increase (decrease) in connection density observed in participants whose perceptual alternation was increased (decreased) by tACS. These results support the hypothesis that deviant visual stimuli evoke θ-band phase synchrony between the frontal and occipital cortices, thereby enhancing perceptual alternation in binocular rivalry.

Theta phase coherence in visual mismatch responses involved in access processing to visual awareness.

Introduction: The electroencephalographic brain response to a deviation from the preceding sequential regularity of visual events, called visual mismatch negativity (vMMN), is well known to reflect automatic visual change detection. Our preliminary study showed a significant correlation between the enhancement of the vMMN amplitude and facilitation of perceptual alternation in binocular rivalry when the deviant stimulus was presented unconsciously. This implies that the vMMN is relevant to access processing, in which the unconscious stimulus is consciously perceived. Recent studies have reported that theta band oscillation evoked by a deviant stimulus is involved in evoking vMMN. However, it has not been clarified whether theta band oscillation associated with vMMN is also relevant to access processing. Methods: We analyzed the correlations between event-related spectral perturbation (ERSP) and inter-trial phase coherence (ITPC) in the theta band and the proportion of perceptual alternation from before to after the presentation of deviation in the same experimental paradigm as in our previous study. Results: We found that an increase in ITPC in the theta band tended to correlate with facilitation of perceptual alternation in binocular rivalry when the deviant was presented unconsciously, but there was no significant correlation in ERSP. Discussion: The results suggest that theta phase coherence underlying the visual mismatch process is relevant to the access processing.

Spatial synaptic modulation through IP3 diffusion triggered by ECB: a computational study with an astrocyte-neurons model.

Recently, functional interactions between neurons and astrocytes have been steadily clarified. In particular, the effects of presynaptic depolarization-induced suppression of excitation (DSE) through endocannabinoid (ECB) and endocannabinoids-mediated synaptic potentiation (eSP) by an astrocyte have been used as an evidence of global heterosynaptic modulation. However, the mechanism of occurrence of spatial modulation in a neural network remains unknown. Although the Ca2+ density in astrocytes is strongly related to eSP through ECB, the mechanism of the rise in the ECB receptor in Ca2+ remains unclear. Since Ca2+ is closely related to inositol-1,4,5-trisphosphate (IP3), it is believed that the released IP3 affects Ca2+ in astrocytes that receive ECB. Therefore, this study approximately showed the spatial distribution of DSE or eSP with astrocyte-neuron computational models, assuming that the IP3 caused by ECB is transmitted in an astrocyte. The results showed doughnut-shaped DSE, eSP, and DSE regions from the central ECB released points to the surroundings. They suggested that IP3 diffusion plays an important role in mediating this synaptic modulation.

Involvement of Visual Mismatch Negativity in Access Processing to Visual Awareness.

Psychophysiological studies with electroencephalography, focusing on the dynamical aspect of neural correlate of consciousness, reported that visual awareness negativity and P3 enhancement are observed at a latency, 200-300 ms after the visual stimulus onset, when the visual stimulus is consciously perceived. However, access processing to visual awareness (APVA) immediately before conscious perception still remains at the earlier stage of visual sensory processing, though there is little known regarding this subject. The present study hypothesized that visual mismatch negativity (vMMN), which reflects automatic change detection at a latency of 130-250 ms, might be involved in the APVA. In a previous study, vMMN was reported to be evoked by the deviant stimulus that is not consciously perceived in binocular rivalry. To clarify whether the visual change detection affects APVA, we conducted a modified experiment of oddball paradigm on binocular rivalry. The results showed a significant correlation between enhancement of vMMN amplitude and facilitation of perceptual alternation when the unconscious deviant was presented. This implies that vMMN is relevant to the APVA, which is a novel role of vMMN. In early visual processing, the attentional mechanism associated with vMMN is suggested to play an important role in unconscious neural processing at an earlier stage of visual awareness.

Early visual processing relevant to the reduction of adaptation-induced perceptual bias.

Visual perception is biased by the preceding visual environment. A well-known perceptual bias is the negative bias where a current percept is biased away from the preceding image (adaptor). The preceding adaptor induces augmentation of early visual evoked potential (the P1 enhancement) of the following test image; the adaptor may invoke certain visual processing for the subsequent test image. However, the visual mechanism underlying P1 enhancement remains unclear. The present study assessed what the P1 alteration reflects in relation to the occurrence of the negative bias. In terms of inter-individual differences, we report that the P1 enhancement of the Necker lattice significantly correlated with the reduction of the reverse-bias effect. Further analyses revealed that the P1 enhancement was insusceptible to neural adaptation to the adaptor at the level of perceptual configuration. The present study suggests that prolonged exposure to a visual image induces modulatory visual processing for the subsequent image (reflected in the P1 enhancement), which is relevant to counteraction of the negative bias.

Cortical dynamics of visual change detection based on sensory memory.

Detecting a visual change was suggested to relate closely to the visual sensory memory formed by visual stimuli before the occurrence of the change, because change detection involves identifying a difference between ongoing and preceding sensory conditions. Previous neuroimaging studies showed that an abrupt visual change activates the middle occipital gyrus (MOG). However, it still remains to be elucidated whether the MOG is related to visual change detection based on sensory memory. Here we tried to settle this issue using a new method of stimulation with blue and red LEDs to emphasize a memory-based change detection process. There were two stimuli, a standard trial stimulus and a deviant trial stimulus. The former was a red light lasting 500 ms, and the latter was a red light lasting 250 ms immediately followed by a blue light lasting 250 ms. Effects of the trial-trial interval, 250 approximately 2000 ms, were investigated to know how cortical responses to the abrupt change (from red to blue) were affected by preceding conditions. The brain response to the deviant trial stimulus was recorded by magnetoencephalography. Results of a multi-dipole analysis showed that the activity in the MOG, peaking at around 150 ms after the change onset, decreased in amplitude as the interval increased, but the earlier activity in BA 17/18 was not affected by the interval. These results suggested that the MOG is an important cortical area relating to the sensory memory-based visual change-detecting system.

Non-linear laws of echoic memory and auditory change detection in humans.

BACKGROUND: The detection of any abrupt change in the environment is important to survival. Since memory of preceding sensory conditions is necessary for detecting changes, such a change-detection system relates closely to the memory system. Here we used an auditory change-related N1 subcomponent (change-N1) of event-related brain potentials to investigate cortical mechanisms underlying change detection and echoic memory. RESULTS: Change-N1 was elicited by a simple paradigm with two tones, a standard followed by a deviant, while subjects watched a silent movie. The amplitude of change-N1 elicited by a fixed sound pressure deviance (70 dB vs. 75 dB) was negatively correlated with the logarithm of the interval between the standard sound and deviant sound (1, 10, 100, or 1000 ms), while positively correlated with the logarithm of the duration of the standard sound (25, 100, 500, or 1000 ms). The amplitude of change-N1 elicited by a deviance in sound pressure, sound frequency, and sound location was correlated with the logarithm of the magnitude of physical differences between the standard and deviant sounds. CONCLUSIONS: The present findings suggest that temporal representation of echoic memory is non-linear and Weber-Fechner law holds for the automatic cortical response to sound changes within a suprathreshold range. Since the present results show that the behavior of echoic memory can be understood through change-N1, change-N1 would be a useful tool to investigate memory systems.

Echoic memory of a single pure tone indexed by change-related brain activity.

BACKGROUND: The rapid detection of sensory change is important to survival. The process should relate closely to memory since it requires that the brain separate a new stimulus from an ongoing background or past event. Given that sensory memory monitors current sensory status and works to pick-up changes in real-time, any change detected by this system should evoke a change-related cortical response. To test this hypothesis, we examined whether the single presentation of a sound is enough to elicit a change-related cortical response, and therefore, shape a memory trace enough to separate a subsequent stimulus. RESULTS: Under a paradigm where two pure sounds 300 ms in duration and 800 or 840 Hz in frequency were presented in a specific order at an even probability, cortical responses to each sound were measured with magnetoencephalograms. Sounds were grouped to five events regardless of their frequency, 1D, 2D, and 3D (a sound preceded by one, two, or three different sounds), and 1S and 2S (a sound preceded by one or two same sounds). Whereas activation in the planum temporale did not differ among events, activation in the superior temporal gyrus (STG) was clearly greater for the different events (1D, 2D, 3D) than the same event (1S and 2S). CONCLUSIONS: One presentation of a sound is enough to shape a memory trace for comparison with a subsequent physically different sound and elicits change-related cortical responses in the STG. The STG works as a real-time sensory gate open to a new event.

A neural network model for exogenous perceptual alternations of the Necker cube.

When a bistable visual image, such as the Necker cube, is continuously viewed, the percept of the image endogenously alternates between one possible percept and the other. However, perceptual alternation can also be induced by an exogenous perturbation. For example, a typical external perturbation is the flashlight, which is expected to pervasively activate many brain regions. Therefore, the neural mechanism related to exogenous perceptual alternation remains to be clarified. As a cue to solving this problem, our recent psychophysiological experiment reported a positive correlation between the enhancement of visual mismatch negativity evoked by breaks in the sequential regularity of the visual stimuli and the proportion of perceptual alternation. To elucidate the mechanism underlying exogenous perceptual alternation induced by visual mismatch negativity, the present study attempted to construct a neural network model for bistable perception of the Necker cube, whose perceptual alternation is facilitated by an increase in visual mismatch negativity. The model consists of both a prediction layer and a prediction error layer, following the predictive coding framework for biologically plausible relationships between the change detection process and the perceptual alternation mechanism. Computer simulations showed that the mean duration of perception decreased as the response increased, which is in concordance with the experimental data. This result suggested that the excitatory feedforward and inhibitory feedback connections play an important role. Additionally, the validity of this model suggests that the visual mismatch signal propagates in the neural systems and affects the visual perceptual mechanism as a prediction error signal.

Correlation Between Gait Perception and Autistic Traits in the General Population: A Study on Event-Related Evoked Potentials.

Previous studies have shown that people with autistic traits have difficulties in motion perception, such as human gait as depicted on a point-light display. A recent study reported that adults with autism spectrum disorders showed atypical visual event-related evoked potentials (ERPs) in response to radial optic flow. To determine the correlation between gait perception and autistic traits in the general population, the present study recorded ERPs time-locked to the onset of approaching and receding point-light walkers. ERPs were measured using an 8-channel system in 19 adults and the correlation between the ERP components and the Subthreshold Autism Trait Questionnaire (SATQ) score were assessed to quantitatively measure autistic traits in the general population. The results showed that the higher SATQ score was, the longer the latency of the ERP component for an approaching walker was. In conclusion, people with autistic traits have trouble perceiving the approach of other people.

Enhancement of a genuine visual mismatch negativity correlates with the facilitation of perceptual alternation of a bistable image.

By focusing on the automatic visual change detection process, the present study attempted to clarify neural processing relevant to the exogenously driven perceptual alternation (ePA) of a bistable image. In our recent electroencephalographic study, the oddball paradigm was adopted to the continuous presentation of a bistable image to record visual mismatch negativity (vMMN, a relative enhancement in the brain response to a deviant over a repetitively presented standard, reflecting the visual change detection process and prediction error to the deviant over the standard). In terms of interindividual differences in behavioral and neural data, a correlation was reported previously between the enhancement of vMMN and facilitation of perceptual alternation, suggesting the involvement of the visual change detection process in ePA. However, the vMMN recorded was expected to be confounded by neural adaptation to the repetitively presented standard; thus, it currently remains unclear whether visual change detection not dependent on neural adaptation, reflected in a 'genuine vMMN', is relevant to ePA. To examine this issue, the present study used a new stimulation paradigm, based on the so-called equiprobable paradigm, to mitigate neural adaptation. The results showed that a genuine vMMN significantly emerged and correlated with an increase in the proportion of perceptual alternation across participants. This supports the involvement of the automatic visual change detection process, not dependent on neural adaptation, in facilitating perceptual alternation. The present results provide a deeper understanding of the involvement of the visual change detection process in ePA.

Exogenously-driven perceptual alternation of a bistable image: From the perspective of the visual change detection process.

Based on the predictive coding framework, the present behavioral study focused on the automatic visual change detection process, which yields a concomitant prediction error, as one of the visual processes relevant to the exogenously-driven perceptual alternation of a bistable image. According to this perspective, we speculated that the automatic visual change detection process with an enhanced prediction error is relevant to the greater induction of exogenously-driven perceptual alternation and attempted to test this hypothesis. A modified version of the oddball paradigm was used based on previous electroencephalographic studies on visual change detection, in which the deviant and standard defined by the bar's orientation were symmetrically presented around a continuously presented Necker cube (a bistable image). By manipulating inter-stimulus intervals and the number of standard repetitions, we set three experimental blocks: HM, IM, and LM blocks, in which the strength of the prediction error to the deviant relative to the standard was expected to gradually decrease in that order. The results obtained showed that the deviant significantly increased perceptual alternation of the Necker cube over that by the standard from before to after the presentation of the deviant. Furthermore, the differential proportion of the deviant relative to the standard significantly decreased from the HM block to the IM and LM blocks. These results are consistent with our hypothesis, supporting the involvement of the automatic visual change detection process in the induction of exogenously-driven perceptual alternation.

Involvement of the visual change detection process in facilitating perceptual alternation in the bistable image.

A bistable image induces one of two perceptual alternatives. When the bistable visual image is continuously viewed, the percept of the image alternates from one possible percept to the other. Perceptual alternation was previously reported to be induced by an exogenous perturbation in the bistable image, and this perturbation was theoretically interpreted to cause neural noise, prompting a transition between two stable perceptual states. However, little is known experimentally about the visual processing of exogenously driven perceptual alternation. Based on the findings of a previous behavioral study (Urakawa et al. in Perception 45:474-482, 2016), the present study hypothesized that the automatic visual change detection process, which is relevant to the detection of a visual change in a sequence of visual events, has an enhancing effect on the induction of perceptual alternation, similar to neural noise. In order to clarify this issue, we developed a novel experimental paradigm in which visual mismatch negativity (vMMN), an electroencephalographic brain response that reflects visual change detection, was evoked while participants continuously viewed the bistable image. In terms of inter-individual differences in neural and behavioral data, we found that enhancements in the peak amplitude of vMMN1, early vMMN at a latency of approximately 150 ms, correlated with increases in the proportion of perceptual alternation across participants. Our results indicate the involvement of automatic visual change detection in the induction of perceptual alternation, similar to neural noise, thereby providing a deeper insight into the neural mechanisms underlying exogenously driven perceptual alternation in the bistable image.

Reduction in the Reverse-Bias Effect by an Abrupt Break in the Sequential Regularity of Visual Events.

A bistable image is more likely to be initially perceived as the reversal of its preceding unambiguous version presented for a prolonged period. This perceptual bias is called the reverse-bias effect. We hypothesized that an abrupt break in the sequential regularity of visual events, synchronized with the onset of a bistable image, counteracts the reverse-bias effect in a similar manner to the disturbing effect of noise in the perceptual process. Under the condition in which the reverse-bias effect was achieved with the Necker lattice, the orientation of the bars around the lattice was simultaneously changed at the onset of the lattice, yielding an abrupt break in the sequential regularity of visual events besides the lattice. The results obtained showed that the reverse-bias effect was significantly reduced by the abrupt break, suggesting that an abrupt break in the sequential regularity of visual events perturbs the perceptual bias of the bistable image, similar to that caused by noise.

Neuromagnetic evidence that the right fusiform face area is essential for human face awareness: An intermittent binocular rivalry study.

When and where the awareness of faces is consciously initiated is unclear. We used magnetoencephalography to probe the brain responses associated with face awareness under intermittent pseudo-rivalry (PR) and binocular rivalry (BR) conditions. The stimuli comprised three pictures: a human face, a monkey face and a house. In the PR condition, we detected the M130 component, which has been minimally characterized in previous research. We obtained a clear recording of the M170 component in the fusiform face area (FFA), and found that this component had an earlier response time to faces compared with other objects. The M170 occurred predominantly in the right hemisphere in both conditions. In the BR condition, the amplitude of the M130 significantly increased in the right hemisphere irrespective of the physical characteristics of the visual stimuli. Conversely, we did not detect the M170 when the face image was suppressed in the BR condition, although this component was clearly present when awareness for the face was initiated. We also found a significant difference in the latency of the M170 (human

Neural substrates of species-dependent visual processing of faces: use of morphed faces.

Face identification and categorization are essential for social communication. The N170 event-related potential (ERP) is considered to be a biomarker of face perception. To elucidate the neural basis of species-dependent face processing, we recorded 128-ch high-density ERPs in 14 healthy adults while they viewed the images of morphed faces. The morphed stimuli contained different proportions of human and monkey faces, and the species boundary was shifted away from the center of the morph continuum. Three experiments were performed to determine how task requirement, facial orientation, and spatial frequency (SF) of visual stimuli affected ERPs. In an equal SF condition, the latency, and amplitude of the occipital P100 for upright faces were modulated in a monotonic-like fashion by the level of morphing. In contrast, the N170 latency for upright faces was modulated in a step-like fashion, showing a flexion point that may reflect species discrimination. Although N170 amplitudes for upright faces were not modulated by morph level, they were modulated in a monotonic-like fashion by inverted faces. The late positive (LP) component (350-550 msec) in the parietal region was modulated in a U-shaped function by morph level during a categorization task, but not in a simple reaction task. These results suggest that P100 reflects changes in the physical properties of faces and that N170 is involved in own-species selectivity. The LP component seems to represent species categorization that occurs 350 msec after stimulus onset.

Proposal for a new MEG-MRI co-registration: a 3D laser scanner system.

OBJECTIVE: Co-registration between the head shape extracted from anatomical images that are obtained using a 3D digitizer is a non-negligible factor for magnetoencephalographic (MEG) utilization. The study aimed to propose a novel quick system based on a laser scanning technique involving a 3D laser scanner system that allows instant measurement while maintaining high accuracy and reproducibility. METHODS: The measurement duration, accuracy, and reproducibility of the finger representations in response to tactile stimulation between the 3D laser scanner-based method and the conventional magnetic field digitizer-based method were compared in 11 healthy subjects. Day-to-day variance in target registration error (TRE), day-to-day and session-to-session variability in head position indicator error (HRE) and source localization accuracy were evaluated with visualization of the source estimation and analysis of variance (ANOVA). RESULTS: Our proposed one-snapshot approach enabled 3D digitization measurements in <5s, and significantly reduced TRE, while mean HREs were a comparable level. Although there was less dramatic improvement of source localization, we found a significant reduction in session-to-session variability and day-to-day variance using our proposed method. CONCLUSIONS: Our results clearly demonstrated improvements in speed, comfort, accuracy, and reproducibility when using our new MEG co-registration method. SIGNIFICANCE: A systematic improvement in MEG measurement will be beneficial for routine clinical use.

Optimum stimulus size for the human brain to respond to motion: a magnetoencephalographic study.

OBJECTIVE: To determine if there is an optimum spatial extent for the detection of moving objects in humans. METHODS: We investigated human brain responses to motion at various speeds (2.9-23.5 deg/s) and stimulus sizes (2.2 × 2.9 deg to 44.8 × 57.4 deg) using magnetoencephalography. The results were compared with those for the flickers with the same stimulus sizes and temporal frequencies. RESULTS: For every size, response latency was inversely related to speed. Further, the latency was lowest at a stimulus size of 16.8 × 22.4 deg for every speed. Although response latency was inversely related to the temporal frequency of the flicker stimulation for all stimulus sizes, it was not affected by stimulus size as much as motion stimulus. CONCLUSIONS: For visual motion detection, the most efficient stimulus size is around 16.8 × 22.4 deg. SIGNIFICANCE: The results suggest that spatial summation mechanism is important for the detection of visual motion but a loss of reference frame information affects the detection of larger motion stimuli, supporting the view that illusory self-motion (vection) is caused by poor reference frame information for motion detection.

Effects of stimulus field size and coherence of visual motion on cortical responses in humans: an MEG study.

Among various kinds of visual motion, wide field coherent visual motion should have characteristic physiological significance regarding the relationship between the external world and us. To detect veridical visual motion in the surrounding environment, specific mechanisms are necessary to differentiate it from the wide field coherent motion due to one's own movement. To disclose whether and how the neural process of wide field coherent motion is different from that of other motions, we measured cortical responses to visual motions in humans using magnetoencephalography (MEG) manipulating both field size and coherence. Results showed that an increase in field size enhanced the response at sensors around the parieto-occipital area, and that the difference in activity between coherent and incoherent motion tended to be larger for the wide field. These findings suggest that wide field coherent and incoherent motion is detected differently at least in part in the parieto-occipital area, and suggest the neural process of wide field coherent motion could be pronouncedly tapped by a combination of field size and coherence.