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Visual working memory (vWM) is an important ability required for various cognitive tasks although its neural underpinnings remain unclear. While many studies have focused on theta (4-7 Hz) and gamma (> 30 Hz) rhythms as a substrate of vWM, here we show that temporal signals embedded in alpha (8-12 Hz) and beta (13-30 Hz) bands can be a good predictor of vWM capacity. Neural activity of healthy human participants was recorded with magnetoencephalography when they performed a classical vWM task (change detection). We analyzed changes in inter-peak intervals (IPIs) of oscillatory signals along with an increase in WM load (a number of to-be-memorized items, 1-6). Results showed a load-dependent reduction of IPIs in the parietal and frontal regions, indicating that alpha/beta rhythms became faster when multiple items were stored in vWM. Furthermore, this reduction in IPIs was positively correlated with individual vWM capacity, especially in the frontal cortex. Those results indicate that vWM is represented as a change in oscillation frequency in the human cerebral cortex., Competing Interests: Declaration of competing interest The authors declare no competing financial interest., (Copyright © 2020. Published by Elsevier Inc.)

Neural oscillatory signals has been associated with many high-level functions (e.g. attention and working memory), because they reflect correlated behaviors of neural population that would facilitate the information transfer in the brain. On the other hand, a decreased power of oscillation (event-related desynchronization, ERD) has been associated with an irregular state in which many neurons behave in an uncorrelated manner. In contrast to this view, here we show that the human ERD is linked to the increased regularity of oscillatory signals. Using magnetoencephalography, we found that presenting a visual stimulus not only induced a decrease in power of alpha (8-12 Hz) to beta (13-30 Hz) rhythms in the contralateral visual cortex but also reduced the mean and variance of their inter-peak intervals (IPIs). This indicates that the suppressed alpha/beta rhythms became faster (reduced mean) and more regular (reduced variance) during visual stimulation. The same changes in IPIs, especially those of beta rhythm, were observed when subjects allocated their attention to a contralateral visual field. Those results revealed a new role of the event-related decrease in alpha/beta power and further suggested that our brain regulates and accelerates a clock for neural computations by actively suppressing the oscillation amplitude in task-relevant regions., (Copyright © 2019 Elsevier Inc. All rights reserved.)

View-invariant face processing emerges early in life. A previous study (Nakato et al., 2009) measured infant hemodynamic responses to faces from the frontal and profile views in the bilateral temporal areas, which have been reported to be involved in face processing using near-infrared spectroscopy. It was reported that 5-month-old infants showed increased oxyhemoglobin (oxy-Hb) responses to frontal faces, but not to profile faces. In contrast, 8-month-old infants displayed increased oxy-Hb responses to profile faces as well as to frontal faces. In this study, we used the experimental method developed in the previous study to investigate the development of view-invariant face processing, every month for 5 months (from the first 3-8 months of life). We longitudinally measured hemodynamic responses to faces from the frontal and profile views in 14 infants. The longitudinal measurements allowed us to investigate individual differences in each participant. We modeled each infant's hemodynamic oxy-Hb responses to frontal and profile faces using linear regression analysis. Processing of profile faces emerged later and underwent larger improvements than that of frontal faces. We also found an anticorrelation between the speed of improvement in face processing and the hemodynamic response to faces at the age of 3- months. Group analysis of the averaged hemodynamic data from the 14 infants using linear regression revealed that the processing of profile faces emerged between 5 and 6 months of age. Infant view-invariant face processing developed first for frontal faces. This was followed by the emergence of processing of profile faces., (Copyright © 2018 The Authors. Published by Elsevier Inc. All rights reserved.)

Previous studies have demonstrated the cortical mechanisms for the gradient of spatial attention in vision and audition, whereas those for touch have yet to be elucidated in detail. In order to examine the within-hand gradient of tactile spatial attention in the cerebral cortex, we used magnetoencephalography (MEG) to record cortical responses to an electrocutaneous stimulation presented randomly to any of the five fingers of the right hand at a random interstimulus interval (750-1250 ms). Participants attended to the index finger, ring finger, or both to detect a rare target stimulus in a sequence of frequent standard stimuli presented to the attended finger(s) by silent counting. Neuromagnetic responses around the contralateral primary and secondary somatosensory cortices (SIc and SIIc) at 50-70 ms and 80-100 ms, respectively, for the stimulation of the index or ring finger were stronger when that finger was attended than when the distant finger was attended or at rest. The amplitude of the SIIc response was also intermediate when the index and ring fingers were simultaneously attended. The SIIc response to the task-irrelevant stimulation of the thumb or little finger increased when the index or ring finger was attended, respectively, suggesting an across-finger gradient of tactile attention. Simultaneous attention to the index and ring fingers decreased the SIIc response to the task-irrelevant stimulation of the intervening middle finger more than that with attention to either one of the two fingers. The earliest attentional sign was observed for the SI M40c response, with the amplitude increasing with the stimulation of the unattended finger. Furthermore, late responses in the temporo-parietal junction (TPJ) and prefrontal cortex (PFC) were stronger with the stimulation of the unattended finger than with that of the attended finger. Thus, the present study provides cortical evidence for the adaptive control of the within-hand, across-finger gradient of tactile attention that depends on whether attention is focused on a single finger or divided into non-adjacent different fingers., (Copyright © 2018 Elsevier Inc. All rights reserved.)

We previously demonstrated that auditory-evoked cortical responses were suppressed by a weak leading stimulus in a manner similar to the prepulse inhibition (PPI) of startle reflexes. The purpose of the present study was to investigate whether a similar phenomenon was present in the somatosensory system, and also whether this suppression reflected an inhibitory process. We recorded somatosensory-evoked magnetic fields following stimulation of the median nerve and evaluated the extent by which they were suppressed by inserting leading stimuli at an intensity of 2.5-, 1.5-, 1.1-, or 0.9-fold the sensory threshold (ST) in healthy participants (Experiment 1). The results obtained demonstrated that activity in the secondary somatosensory cortex in the hemisphere contralateral to the stimulated side (cSII) was significantly suppressed by a weak leading stimulus with the intensity larger than 1.1-fold ST. This result implied that the somatosensory system had an inhibitory process similar to that of PPI. We then presented two successive leading stimuli before the test stimulus, and compared the extent of suppression between the test stimulus-evoked responses and those obtained with the second prepulse alone and with two prepulses (first and second) (Experiment 2). When two prepulses were preceded, cSII responses to the second prepulse were suppressed by the first prepulse, whereas the ability of the second prepulse to suppress the test stimulus remained unchanged. These results suggested the presence of at least two individual pathways; response-generating and inhibitory pathways., (Copyright © 2014 Elsevier Inc. All rights reserved.)

Grapheme-color synesthesia is a phenomenon in which achromatic letters/digits automatically induce particular colors. When multiple letters are integrated into a word, some synesthetes perceive that all those letters are changed into the same color, reporting lexical color to that word. Previous psychological studies found several "rules" that determine those lexical colors. The colors to most words are determined by the first letters of the words, while some words in ordinal sequences have their specific colors. Recent studies further reported the third case where lexical colors might be influenced by semantic information of words. Although neural mechanisms determining those lexical colors remained unknown, here we identified three separate neural systems in the synesthete's brain underlying three rules for illusory coloring of words. In addition to the occipito-temporal and parietal regions previously found to be associated with the grapheme-color synesthesia, neural systems for lexical coloring extended to linguistic areas in the left inferior frontal and anterior temporal regions that were engaged in semantic analyses of words. Those results indicate an involvement of wider and higher neural networks than previously assumed in a production of synesthetic colors to visual stimuli and further showed a multiplicity of synesthetic mechanisms represented in the single brain., (Copyright © 2014 Elsevier Inc. All rights reserved.)

Changes in continuous sounds elicit a preattentive component that peaks at around 100ms (Change-N1m) on electroencephalograms or magnetoencephalograms (MEG). Change-N1m is thought to reflect brain activity relating to the automatic detection of changes, which facilitate processes for the execution of appropriate behavior in response to new environmental events. The aim of the present MEG study was to elucidate whether a component relating to auditory changes existed earlier than N1m. Change-related cortical responses were evoked by abrupt sound movement in a train of clicks at 100Hz. Sound movement was created by inserting an interaural time delay (ITD) of 0.15, 0.25, 0.35, and 0.45ms into the right ear. Ten out of 12 participants exhibited clear change-related cortical responses earlier than Change-N1m at around 60ms (Change-P50m). The results of source analysis showed that Change-P50m originated from the superior temporal gyrus of both hemispheres and that its location did not differ significantly from dipoles for the response to the sound onset. The magnitude of Change-P50m increased and the peak latency shortened with an increase in the ITD, similar to those of Change-N1m. These results suggest that change-related cortical activity is present as early as its onset latency at around 50ms., (© 2013 Elsevier Inc. All rights reserved.)

This study investigated how infants perceive and interpret human body movement. We recorded the eye movements and pupil sizes of 9- and 12-month-old infants and of adults (N=14 per group) as they observed animation clips of biomechanically possible and impossible arm movements performed by a human and by a humanoid robot. Both 12-month-old infants and adults spent more time looking at the elbows during impossible compared with possible arm movements, irrespective of the appearance of the actor. These results suggest that by 12 months of age, infants recognize biomechanical constraints on how arms move, and they extend this knowledge to humanoid robots. Adults exhibited more pupil dilation in response to the human's impossible arm movements compared with the possible ones, but 9- and 12-month-old infants showed no differential pupil dilation to the same actions. This finding suggests that the processing of human body movements might still be immature in 12-month-olds, as they did not show an emotional response to biomechanically impossible body movements. We discuss these findings in relation to the hypothesis that perception of others' body movements relies upon the infant's own sensorimotor experience., (Copyright © 2012 Elsevier Inc. All rights reserved.)

The role of sensory information in motor control has been studied, but the cortical processing underlying cross-modal relationship between visual and somatosensory information for movement execution remains a matter of debate. Visual estimates of limb positions are congruent with proprioceptive estimates under normal visual conditions, but a mismatch between the watched and felt movement of the hand disrupts motor execution. We investigated whether activation in somatosensory areas was affected by the discordance between the intended and an executed action. Subjects performed self-paced thumb movement of the left hand under normal visual and mirror conditions. The Mirror condition provided a non-veridical and unexpected visual feedback. The results showed activity in the primary somatosensory area to be inhibited and activity in the secondary somatosensory area (SII) to be enhanced with voluntary movement, and neural responses in the SII and parietal cortex were strongly affected by the unexpected visual feedback. These results provide evidence that the visual information plays a crucial role in activation in somatosensory areas during motor execution. A mechanism that monitors sensory inputs and motor outputs congruent with current intension is necessary to control voluntary movement., (Copyright © 2011. Published by Elsevier Inc.)

Arcimboldo images induce the perception of faces when shown upright despite the fact that only nonfacial objects such as vegetables and fruits are painted. In the current study, we examined whether infants recognize a face in the Arcimboldo images by using the preferential looking technique and near-infrared spectroscopy (NIRS). In the first experiment, we measured looking preference between upright and inverted Arcimboldo images among 5- and 6-month-olds and 7- and 8-month-olds. We hypothesized that if infants perceive the Arcimboldo images as faces, they would prefer the upright images to the inverted ones. We found that only 7- and 8-month-olds significantly preferred upright images, suggesting that they could perceive the Arcimboldo images as faces. In the second experiment, we measured hemodynamic responses using NIRS. Based on the behavioral data, we hypothesized that 7- and 8-month-olds would show different neural activity for upright and inverted Arcimboldo images, as do adults. Therefore, we measured hemodynamic responses in 7- and 8-month-olds while they were looking at upright and inverted Arcimboldo images. Their responses were then compared with the baseline activation during the presentation of individual vegetables. We found that the concentration of oxyhemoglobin increased in the left temporal area during the presentation of the upright images compared with the baseline during the presentation of vegetables. The results of the two experiments suggest that (a) the ability to recognize the upright Arcimboldo images as faces develops at around 7 or 8 months of age and (b) processing of the upright Arcimboldo images is related to the left temporal area of the brain., (Copyright © 2011 Elsevier Inc. All rights reserved.)

Interactive non-linear cooperation of different feature dimensions, feature synergy, has been studied in psychophysics, but the neural mechanism is unknown. The present study investigated the neural representation of feature synergy of two second-order visual features by combining electroencephalography (EEG) with the signal detection theory (SDT). Two kinds of a 27-by-27 array of Gabor patches were presented in a random order; a reference stimulus which has no segregated region, and a target stimulus whose inner region differed in spatial frequency, orientation, or both from the surround. Subjects performed a Yes-No discrimination of whether the inner region was different from the surround, while EEG signals were recorded from 62 locations. When the SDT measure showed feature synergy, EEG activity showed a long-lasting enhancement starting at 130 ms around the inferior temporal region. In contrast, no EEG modulation was observed when feature synergy was not present. Thus, our combined approach demonstrates that non-linear cooperation between different features is represented by neural activity starting at 130 ms post-stimulus in the ventral visual stream., (Copyright © 2010 Elsevier Inc. All rights reserved.)

Recognition of other people's facial expressions of emotion plays an important role in social communication in infants as well as adults. Evidence from behavioral studies has demonstrated that the ability to recognize facial expressions develops by 6 to 7 months of age. Although the regions of the infant brain involved in processing facial expressions have not been investigated, neuroimaging studies in adults have revealed that several areas including the superior temporal sulcus (STS) participate in the processing of facial expressions. To examine whether the temporal area overlying the STS is responsible for the processing of facial expressions in infants, near-infrared spectroscopy (NIRS) was used to measure the neural activity in the temporal area overlying the STS as infants looked at happy and angry faces. NIRS provides a non-invasive means of estimating cerebral blood flow in the human brain and does not require severe constraints of head-movement. According to the International 10-20 system for EEG electrode placement, the measurement area was located in the bilateral temporal area centered at positions T5 and T6, which correspond to the STS. The time-course of the average change in total-Hb concentration revealed a clear difference in the pattern of hemodynamic responses to happy and angry faces. The hemodynamic response increased gradually when infants looked at happy faces and was activated continuously even after the disappearance of the face. In contrast, the hemodynamic responses for angry faces increased during the presentation of angry faces, then decreased rapidly after the face disappeared. Moreover, the left temporal area was significantly activated relative to the baseline when infants looked at happy faces, while the right temporal area was significantly activated for angry faces. These findings suggest hemispheric differences in temporal areas during the processing of positive and negative facial expressions in infants., (Copyright © 2010 Elsevier Inc. All rights reserved.)

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., (Copyright 2010 Elsevier Inc. All rights reserved.)

The quick detection of dynamic changes in multisensory environments is essential to survive dangerous events and orient attention to informative events. Previous studies have identified multimodal cortical areas activated by changes of visual, auditory, and tactile stimuli. In the present study, we used magnetoencephalography (MEG) to examine time-varying cortical processes responsive to unexpected unimodal changes during continuous multisensory stimulation. The results showed that there were change-driven cortical responses in multimodal areas, such as the temporo-parietal junction and middle and inferior frontal gyri, regardless of the sensory modalities where the change occurred. These multimodal activations accompanied unimodal activations, both of which in general had some peaks within 300 ms after the changes. Thus, neural processes responsive to unimodal changes in the multisensory environment are distributed at different timing in these cortical areas.

In the present study, we investigate the top-down centrifugal modulation of neural responses to a task-relevant noxious stimulation triggering voluntary movement by recording magnetoencephalography (MEG) and electroencephalography (EEG) simultaneously. An auditory warning signal was followed 2-3 s later by a noxious YAG laser stimulation as an imperative signal delivered to the left hand dorsum. Ten normal subjects performed three different conditions, Control, Movement, and Count. In Control, the subjects were asked to relax and rest quietly with no task. In Movement, the subjects extended the left index finger after imperative stimuli. In Count, the subjects counted the number of imperative stimuli silently. The amplitude of the N2 component recorded by EEG, which peaked about 220 ms after noxious stimulation, was significantly attenuated in Movement, but not in Count, compared to Control. The root-mean-square (RMS) from both hemispheres, and areal mean signal (AMS) amplitudes and the equivalent current dipole (ECD) strengths from SI/PPC and bilateral SII recorded at around 170 ms by MEG were not significantly different among the three conditions. In contrast, ECD strengths and AMS amplitudes from the anterior cingulate cortex (ACC), which showed a similar peak to the N2 component, were smaller in Movement than Control and Count. We therefore suspect that neural activities related to generator mechanisms of N2, especially including ACC, are inhibited by movement-related neural activities during the preparatory period. The present findings indicate a characteristic of pain-motor integration in a movement preparatory period.

We recorded cortical activities in response to the onset and offset of a train of electrical pulses applied to the right hand in eleven healthy volunteers by the use of magnetoencephalograms to clarify temporal and spatial profiles of the somatosensory on- and off-cortical responses. Results showed that a region around the upper bank of the sylvian fissure of both hemispheres responded to the onset and offset of the stimulus, while the activity in the primary somatosensory cortex (SI) of the hemisphere contralateral to the stimulation was clear only for the onset response. The SI activity consisted of two components suggesting that two distinct populations of neurons in SI were involved in processing a train of pulses. The location of the source of activity in the contralateral para-sylvian region (cPara) differed significantly between the on- and off-response, while that of the activity in the ipsilateral para-sylvian region (iPara) did not. The differences in location of the cPara activity might be caused by the overlapping of several cortical activities in response to each stimulus and stimulus event (on and off events), while the iPara activity might reflect purely the event-related response. Moreover, some subjects had clear iPara activity without cPara activity especially in the off-response, suggesting the iPara activity to be independent of the cPara activity. We consider that activities in the parasylvian region are involved in the detection of changes at the body's surface.

Primary somatosensory cortex (SI) and posterior parietal cortex (PPC) are activated by noxious stimulation. In neurophysiological studies using magnetoencephalography (MEG), however, it has been difficult to separate the activity in SI from that in PPC following stimulation of the upper limb, since the hand area of SI is very close to PPC. Therefore, we investigated human pain processing using MEG following the application of a thulium-YAG laser to the left thigh to separate the activation of SI and PPC, and to clarify the time course of the activities involved. The results indicated that cortical activities were recorded around SI, contralateral secondary somatosensory cortex (cSII), ipsilateral secondary somatosensory cortex (iSII), and PPC between 150-185 ms. The precise location of PPC was indicated to be the inferior parietal lobule (IPL), corresponding to Brodmann's area 40. The mean peak latencies of SI, cSII, iSII and IPL were 152, 170, 181, and 183 ms, respectively. This is the first study to clarify the time course of the activities of SI, SII, and PPC in human pain processing using MEG.

This is the first functional magnetic resonance imaging (fMRI) study to investigate the hemodynamic response related to somatosensory spatial discrimination, so-called two-point discrimination. During scanning, we examined two discrimination tasks using four types of electrical stimuli applied to one or two points with strong or weak intensity on the right and left forearm, respectively. In the two-point discrimination task (TPD), subjects reported whether they thought the stimulus was applied to one point or two. In the intensity discrimination task (ID), subjects were required to judge whether the stimulus was strong or weak. In each task, they pressed a button to report their choice. Comparing TPD with the control, we found activated regions in the inferior parietal lobule (IPL) around the supramarginal gyrus (SMG) (Brodmann's area 40) and anterior cingulate cortex (ACC). These areas were significantly activated irrespective of the forearm stimulated. Comparing ID with the control, there were no significantly activated regions. By comparing the TPD and ID, we identified that the left IPL was significantly activated, specifically in TPD, irrespective of the forearm stimulated. In contrast, there were no significantly activated regions in the ID task. Therefore, the left IPL is considered to play an important role in two-point discrimination.

Inhibiting inappropriate behavior and thoughts is an essential ability for humans, but the regions responsible for inhibitory processing are a matter of continuous debate. This is the first study of somatosensory go/nogo tasks using event-related functional magnetic resonance imaging (fMRI). Fifteen subjects preformed two different types of go/nogo task, i.e. (1) Movement and (2) Count, to compare with previous studies using visual go/nogo tasks, and confirm whether the inhibitory processing is dependent on sensory modalities. Go and nogo stimuli were presented with an even probability. Our data indicated that the response inhibition network involved the dorsolateral (DLPFC) and ventrolateral (VLPFC) prefrontal cortices, pre-supplementary motor area (pre-SMA), anterior cingulate cortex (ACC), inferior parietal lobule (IPL), insula, and temporoparietal junction (TPJ), which were consistent with previous results obtained using visual go/nogo tasks. These activities existed in both Movement and Count Nogo trials. Therefore, our results suggest that the network for inhibitory processing is not dependent on sensory modalities but reflects common neural activities. In addition, there were differences of activation intensity between Movement and Count Nogo trials in the prefrontal cortex, temporal lobe, and ACC. Thus, inhibitory processing would involve two neural networks, common and uncommon regions, depending on the required response mode.

We investigated whether early activity in the occipitotemporal region, corresponding to human MT/V5, is influenced by a face contour and/or features such as the mouth using magnetoencephalography (MEG). We used apparent motion as visual stimuli and compared four conditions as follows: (1) CDL: A schematic face consisting of a face Contour, two Dots and a horizontal Line (2) CD: The Contour and two Dots (3) DL: Two Dots and a horizontal Line and (4) D: Two Dots only. Subjects described a simple movement of dots for D, but eye movement for CDL, DL and CD, though movement modalities were the same through all conditions. We used a single equivalent current dipole (ECD) model within 145-220 ms after stimulus onset and estimated the location, dipole moment (strength) and peak latency. There were no significant differences in the peak latency of the estimated dipoles between each condition, but the activity was significantly stronger for CDL than for CD (p<0.05), DL (p<0.01), and D (p<0.01) in the right hemisphere, and DL and D (p<0.01) in the left. These results indicated that there is specific information processing for eye movements in the occipitotemporal region, the human MT/V5 homologue, and this activity was significantly influenced by whether movements appeared with the face contour and/or features, in other words, whether the eyes moved or not, even if the movement itself was the same.

To investigate the motor control related to sound production, we studied cortical rhythmic changes during continuous vocalization such as singing. Magnetoencephalographic (MEG) responses were recorded while subjects spoke in the usual way (speaking), sang (singing), hummed (humming) and imagined (imagining) a popular song. The power of alpha (8-15 Hz), beta (15-30 Hz) and low-gamma (30-60 Hz) frequency bands was changed during and after vocalization (singing, speaking and humming). In the alpha band, the oscillatory changes for singing were most pronounced in the right premotor, bilateral sensorimotor, right secondary somatosensory and bilateral superior parietal areas. The beta oscillation for the singing was also confirmed in the premotor, primary and secondary sensorimotor and superior parietal areas in the left and right hemispheres where were partly activated even for imagined a song (imaging). These regions have been traditionally described as vocalization-related sites. The cortical rhythmic changes were distinct in the singing condition compared with the other vocalizing conditions (speaking and humming) and thus we considered that more concentrated control of the vocal tract, diaphragm and abdominal muscles is responsible. Furthermore, characteristic oscillation in the high-gamma (60-200 Hz) frequency band was found in Broca's area only in the imaging condition and might occur singing rehearsal and storage process in Broca's area.

The present study examined infants' brain activity in response to upright and inverted faces using near infrared spectroscopy (NIRS), which can non-invasively record hemodynamic changes of the brain. NIRS is particularly useful for recording in infants, since recordings can be made, even while the infants are awake, without fixing their body and brain. For this objective, we used newly developed sensor probes of NIRS for recording in infants. We measured changes in cerebral oxygenation in 10 5-8-month-olds' left and right lateral areas while they were looking at upright and inverted faces. The results are summarized as follows: (1) the concentration of oxyhemoglobin (oxy-Hb) and total hemoglobin (total-Hb) increased significantly in the right lateral area during the upright face condition, (2) the concentration of total-Hb in the right lateral area differed significantly between the upright and inverted conditions, (3) hemodynamic changes were maximal in the temporal region, probably in the superior temporal sulcus (STS) in both hemispheres, and (4) the right hemisphere seems to be more important for recognizing upright faces. This is the first evidence showing that there is an inter-hemispheric difference on the effect of face inversion in the infant brain using a hemodynamic method.

Many studies have reported a movement-related modulation of response in the primary and secondary somatosensory cortices (SI and SII) to a task-irrelevant stimulation in primates. In the present study, magnetoencephalography (MEG) was used to examine the top-down centrifugal regulation of neural responses in the human SI and SII to a task-relevant somatosensory signal triggering a voluntary movement. Nine healthy adults participated in the study. A visual warning signal was followed 2 s later by a somatosensory imperative signal delivered to the right median nerve at the wrist. Three kinds of warning signal informed the participants of the reaction which should be executed on presentation of the imperative signal (rest or extension of the right index finger, extension of the left index finger). The somatosensory stimulation was used to both generate neural responses and trigger voluntary movement and therefore was regarded as a task-relevant signal. The responses were recorded using a whole-head MEG system. The P35m response around the SI was reduced in magnitude without alteration of the primary SI response, N20m, when the signal triggered a voluntary movement compared to the control condition, whereas bilateral SII responses peaking at 70-100 ms were enhanced and the peak latency was shortened. The peak latency of the responses in the SI and SII preceded the onset of the earliest voluntary muscle activation in each subject. Later bilateral perisylvian responses were also enhanced with movement. In conclusion, neural activities in the SI and SII evoked by task-relevant somatosensory signals are regulated differently by motor-related neural activities before the afferent inputs. The present findings indicate a difference in function between the SI and SII in somatosensory-motor regulation.

To clarify the latency of the earliest cortical activity in visual processing, electroretinograms (ERGs) and visual evoked magnetic fields (VEFs) following flash stimulation were recorded simultaneously in six human subjects. Flash stimuli were applied to the right eye and ERGs were recorded from a skin electrode placed on the lower lid. ERGs showed two major deflections in all subjects: an eyelid-negativity around 20 ms and a positivity around 60 ms corresponding to an a- and b-waves, respectively. The mean onset and peak latency of the earliest component of VEFs (37 M) was 30.2 and 36.9 ms, respectively. There was a linear correlation between the peak latency of the a-wave and the onset latency of the 37 M (r=0.90, P=0.011). When a single equivalent current dipole analysis was applied to the 37 M, four out of six subjects showed highly reliable results. The generator of the 37 M was estimated to be located in the striate cortex in all four subjects. Since post-receptoral activities in the retina are expected to start around the peak of the a-wave (20 ms), the early cortical activity, which appears 10 ms later than the a-wave peak, is considered to be the earliest cortical activity following flash stimulation.

The objective of this study was to reveal the oscillatory activity in the occipitotemporal area related to the visual perception of the letters of first (L1) and second languages (L2) and pseudoletters. We recorded neuromagnetic signals while Korean native speakers were exposed to a phonogram of Korean, acquired at school age as their L1 (Hangul), that of Japanese, learned in adulthood as a L2 (Kana) and pseudoletter (Pseudo), and quantified the event-related desynchronization (ERD) and synchronization (ERS). In all conditions, sustained ERDs in the alpha band were observed in both hemispheres. ERD for Pseudo was gradually attenuated after approximately 400-500 ms after stimulus onset, whereas both Hangul and Kana produced stronger and longer-lasting ERD. ERD for Kana showed a broader alpha band than Hangul. Furthermore, transient ERSs in the gamma band around 70 Hz were observed between 100 and 400 ms in the bilateral occipitotemporal areas. In the left hemisphere, gamma band oscillations showed similar enhancement in all conditions, suggesting that gamma band activity in the left occipitotemporal area might be enhanced not only by the bottom-up process as visual perception but also by the top-down process as attention to prelexical visual stimuli. In the right hemisphere, gamma band ERS was stronger for Hangul than Pseudo and no differences were shown between Kana and Pseudo. The differences of oscillatory activity in the alpha and gamma bands suggest that neuronal networks, including the occipitotemporal area, are related to the visual perception of letters differing between L1 and L2.

Backward masking is one of the potent ways to reveal the neural mechanism of visual awareness in humans. Although previous neuroimaging studies have reported that the visual masking involves the attenuation of hemodynamic signals to the masked stimulus in visual ventral regions such as the fusiform and inferior temporal gyrus, the temporal profiles of this attenuation as a whole neural population is mostly unclear. Here we used magnetoencephalography and investigated the neural response changes in higher visual region induced by backward masking. The combination of our previous random dot blinking method with the sensor-based analysis isolated the neural responses in the higher visual cortex relating to shape perception. The results revealed that, as the visibility of the target stimulus was reduced by the mask following it, the neural response to the target in the ventral regions showed gradual decreases both in its peak amplitude and peak latency. Furthermore, this decrease in the peak amplitudes was significantly correlated with the behavioral accuracy of the target identification, while the peak latency was not. These results indicate that backward masking simultaneously produces two types of neural changes in higher visual regions: attenuation of the populational neural activity itself and temporal interruption of this activity by the subsequent mask response. Especially, our data suggest that the response attenuation in higher visual response is a main cause of the perceptual impairment observed in the backward masking paradigm.

The temporal change in somatosensory evoked magnetic fields (SEFs) in the preparatory period of self-initiated voluntary movement was investigated. The SEF following stimulation of the right median nerve was recorded, using a 204-channel whole-head MEG system, in nine healthy subjects during a self-initiated extension of the right index finger every 5 to 7 s. The preparatory period before finger movement was divided into six subperiods, and the MEG signals following the stimulation in each subperiod were averaged separately. SEFs were also recorded in the resting state. The ECD strengths for N20m and P60m were not significantly changed in any subperiod before movement compared with those in the resting state. The ECD strength for P30m was significantly smaller 500 ms or less before movement than during the resting state and 1,500 ms or less before movement compared to that during the period from 3,000 to 4,000 ms before movement. Thus, we confirmed that the SEF components were attenuated even during a period of self-initiated voluntary movement. The modulation started at least 1,500 ms before movement and was greater for the P30m than the N20m component. These findings suggested that motor-associated cortices attenuated SEF components by a centrifugal gating process.

We examined the influence of stimulus duration of foreign consonant vowel stimuli on the MMNm (magnetic counter part of mismatch negativity). In Experiment 1, /ra/ and /la/ stimuli were synthesized and subjects were native Japanese speakers who are known to have difficulty discriminating the stimuli. "Short" duration stimuli were terminated in the middle of the consonant-to-vowel transition (110 ms). They were nevertheless clearly identifiable by English speakers. A clear MMNm was observed only for short-duration stimuli but not for untruncated long-duration (150-ms) stimuli. We suggest that the diminished MMNm for longer duration stimuli result from more effective masking by the longer vowel part. In Experiment 2 we examined this hypothesis by presenting only the third formant (F3) component of the original stimuli, since the acoustic difference between /la/ and /ra/ is most evident in the third formant, whereas F1 and F2 play a major role in vowel perception. If the MMNm effect depends on the acoustic property of F3, a stimulus duration effect comparable to that found with the original /la/ and /ra/ stimuli might be expected. However, if the effect is attributable to the masking effect from the vowel, no influence of stimulus duration would be expected, since neither stimulus contains F1 and F2 components. In fact, the results showed that the "F3 only" stimuli did not show a duration effect; MMNm was always elicited independent of stimulus duration. The MMN stimulus duration effect is thus suggested to come from the backward masking of foreign consonants by subsequent vowels.

Our previous study (T. Nihashi et al., 2001, Neuro- Image 13: 295-304), using magnetoencephalography (MEG), revealed somatotopy of the ear in the human primary somatosensory cortex (SI); that is, the signals following stimulation of the ear reach both the neck and face areas of the SI. However, since this was the first report on somatotopy of the ear in humans, we performed an fMRI activation study to confirm the somatotopic representation, and compared the electrical activity by MEG and the cerebral blood flow change by fMRI. We studied eight healthy subjects using 3-T MRI. We stimulated three parts of the left ear: the helix, the lobulus, and the tragus. First, we identified the location of the ear area in the SI based on our previous MEG study, in which equivalent current dipoles (ECDs) were located in the neck and/or face areas of the SI. Then, we determined the search volume as a sphere with a 15-mm radius, which was placed in the neck and/or face area. We analyzed whether or not fMRI activation occurred inside such spheres. Stimulation of the helix activated the neck area of the SI in four of eight subjects, and both the neck and face areas in two. No activation was observed in two subjects. Stimulation of the lobulus activated the neck area in one subject, the face area in two, both in four, and neither in one. Stimulation of the tragus activated the face in four, both in three, and neither in one. These fMRI findings confirm the result of MEG that the representation of the ear in the SI is separated into neck and face areas.

Stimulus repetition improves performance and modulates event-related brain potentials in word recognition tasks. We recorded evoked magnetic responses from bilateral temporal sites of the brain to determine the cortical area related to the word repetition effect. Fourteen Japanese volunteers read words or pronounceable nonwords, some of which occurred twice with a lag of eight items. Clear magnetic responses were observed bilaterally. In the left hemisphere, a reduction of the magnetic responses by repetition was observed for words but not for nonwords in the latency range of 300-500 ms poststimulus. The sources of the responses were estimated to be in the left perisylvian area adjacent to the auditory cortex and the left parietal area. Only the perisylvian source activity showed the reduction by the word repetition. The left perisylvian area was thus suggested to be related to the word repetition effect. The activity in this area might be associated with the lexical memory process.

The temporal and spatial processing of viewing eye movements was studied by magnetoencephalography (MEG) in six normal subjects. Three visual stimulus types were studied: (1) moving eyes (EYES), (2) moving simulated eyes (SIM), consisting of checks moving in the same spatial location as EYES, and (3) an inwardly moving radial pattern (RADIAL). A large clear MEG component, 1M, with mean peak latency of approximately 170 ms, was seen in the right hemisphere to RADIAL and EYES in all six subjects. The 1M to EYES was significantly longer in latency and smaller in amplitude than that seen to RADIAL. A left hemisphere 1M to EYES and RADIAL was seen in three of six subjects. In all subjects and both hemispheres the equivalent current dipoles (ECD) for EYES and RADIAL were located near the occipitotemporal border, the MT/V5 homologue in humans. The ECD to EYES was significantly more posterior and inferior than that to RADIAL, with a calculated significant separation distance of around 1 cm. No ECD was estimated in the fusiform gyrus, a structure that plays a main role in static face perception. Although the 1M was detected in SIM in all six subjects, our criteria for a reliable ECD could only be satisfied in only one subject. Our results suggest that the cortex of human MT/V5 and its surrounds is active both in the perception of eye motion and motion in general, particularly in the right hemisphere. The areas responsive to eye motion were separable from those responsive to radial motion. These data suggest that there may be specialization within regions of human cortex previously thought to be sensitive to motion in general., (Copyright 2001 Academic Press.)

We studied 13 healthy subjects with a multichannel magnetoencephalography (MEG) system to investigate the somatotopic representation of the ear in the primary somatosensory cortex (SI). We stimulated three parts of the left ear: the helix, the lobulus, and the tragus. The somatosensory-evoked magnetic fields (SEFs) were successfully measured in 7 of 13 subjects. Short-latency responses were analyzed using both single dipole and multidipole models (brain electric source analysis, BESA). From the single dipole model, the equivalent current dipole (ECD) following the helix stimulation was estimated to be near the neck area of SI in all the subjects. In the lobulus stimulation, the ECDs were estimated around the neck area of SI in four subjects, in the face area in one subject, and in the deep white matter in two subjects. In the tragus stimulation, the ECDs were estimated around the neck area of SI in three subjects, in the hand area of SI in two subjects, and in the deep white matter in two subjects. When the ECDs were estimated to be located in unlikely sites (hand area and deep white matter), a two-dipole model, (1) the neck area of SI and (2) face area of SI, was found to be the most appropriate. Although this might be a preliminary study due to a relatively small number of subjects, it revealed that receptive fields of some part of the ear, such as the lobulus and tragus, might be present in both the neck and face areas of SI. These findings suggested that the "ear area" of SI has variability between subjects, unlike the other areas of SI, possibly because the ear is located on the border between the neck and face., (Copyright 2001 Academic Press.)

We studied a detailed somatosensory representation map of the human primary somatosensory cortex using magnetoencephalography. Somatosensory-evoked magnetic fields following tactile stimulation of multiple points in the right hemibody (including the tongue, lips, fingers, arm, trunk, leg, and foot) were analyzed in five normal subjects. We were able to estimate equivalent current dipoles (ECDs) following stimulation of the tongue, lips, fingers, palm, forearm, elbow, upper arm, and toes in most subjects and those following the stimulation of the chest, ankle, and thigh in one subject. The ECDs were located in the postcentral gyrus and generally arranged in order along the central sulcus, which is compatible with the somatosensory "homunculus." Linear distances, averaged in five subjects, from the receptive area of the thumb to that of the tongue, little finger, forearm, upper arm, and toes were estimated to be 2.42 +/- 0.28, 1.25 +/- 0.28, 2.21 +/- 0.72, 2.75 +/- 0.63, and 5.29 +/- 0.48 cm, respectively. The moment of each ECD, which suggested the size of the cortical areas responsive to the stimulation, was also compatible with the bizarre proportion of the homunculus with a large tongue, lips, and fingers. According to these results, we were able to reproduce a large part of the somatosensory homunculus quantitatively on an individual brain MRI.