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Haptic memory briefly retains somatosensory information for later use; however, how and which cortical areas are affected by haptic memory remain unclear. We used change-related cortical responses to investigate the relationship between the somatosensory cortex and haptic memory objectively. Electrical pulses, at 50 Hz with a duration of 500 ms, were randomly applied to the second, third, and fourth fingers of the right and left hands at an even probability every 800 ms. Each stimulus was labeled as D (preceded by a different side) or S (preceded by the same side). The D stimuli were further classified into 1D, 2D, and 3D, according to the number of different preceding stimuli. The S stimuli were similarly divided into 1S and 2S. The somatosensory-evoked magnetic fields obtained were divided into four components via a dipole analysis, and each component's amplitudes were measured using the source strength waveform. The results showed that the preceding event did not affect the amplitude of the earliest 20-30 ms response in the primary somatosensory cortex. However, in the subsequent three components, the cortical activity amplitude was largest in 3D, followed by 2D, 1D, and S. These results indicate that such modulatory effects occurred somewhere in the somatosensory processing pathway higher than Brodmann's area 3b. To the best of our knowledge, this is the first study to demonstrate the existence of haptic memory for somatosensory laterality and its impact on the somatosensory cortex using change-related cortical responses without contamination from peripheral effects., (© 2020 The Authors. Human Brain Mapping published by Wiley Periodicals LLC.)

Paired-pulse suppression refers to attenuation of neural activity in response to a second stimulus and has a pivotal role in inhibition of redundant sensory inputs. Previous studies have suggested that cortical responses to a somatosensory stimulus are modulated not only by a preceding same stimulus, but also by stimulus from a different submodality. Using magnetoencephalography, we examined somatosensory suppression induced by three different conditioning stimuli. The test stimulus was a train of electrical pulses to the dorsum of the left hand at 100 Hz lasting 1500 ms. For the pulse train, the intensity of the stimulus was abruptly increased at 1200 ms. Cortical responses to the abrupt intensity change were recorded and used as the test response. Conditioning stimuli were presented at 600 ms as pure tones, either innocuous or noxious electrical stimulation to the right foot. Four stimulus conditions were used: (1) Test alone, (2) Test + auditory stimulus, (3) Test + somatosensory stimulus, and (4) Test + nociceptive stimulus. Our results showed that the amplitude of the test response was significantly smaller for conditions (3) and (4) in the secondary somatosensory cortex contralateral (cSII) and ipsilateral (iSII) to the stimulated side as compared to the response to condition (1), whereas the amplitude of the response in the primary somatosensory cortex did not differ among the conditions. The auditory stimulus did not have effects on somatosensory change-related response. These findings show that somatosensory suppression was induced by not only a conditioning stimulus of the same somatosensory submodality and the same cutaneous site to the test stimulus, but also by that of a different submodality in a remote area.

Cross-modal interaction occurs during the early stages of processing in the sensory cortex; however, its effect on neuronal activity speed remains unclear. We used magnetoencephalography to investigate whether auditory stimulation influences the initial cortical activity in the primary somatosensory cortex. A 25-ms pure tone was randomly presented to the left or right side of healthy volunteers at 1000 ms when electrical pulses were applied to the left or right median nerve at 20 Hz for 1500 ms because we did not observe any cross-modal effect elicited by a single pulse. The latency of N20 m originating from Brodmann's area 3b was measured for each pulse. The auditory stimulation significantly shortened the N20 m latency at 1050 and 1100 ms. This reduction in N20 m latency was identical for the ipsilateral and contralateral sounds for both latency points. Therefore, somatosensory-auditory interaction, such as input to the area 3b from the thalamus, occurred during the early stages of synaptic transmission. Auditory information that converged on the somatosensory system was considered to have arisen from the early stages of the feedforward pathway. Acceleration of information processing through the cross-modal interaction seemed to be partly due to faster processing in the sensory cortex.

Sensory suppression is a mechanism that attenuates selective information. As for long-latency suppression in auditory and somatosensory systems, paired-pulse suppression, observed as 2 identical stimuli spaced by approximately 500 ms, is widely known, though its mechanism remains to be elucidated. In the present study, we investigated the relationship between auditory and somatosensory long-latency suppression of change-related cortical responses using magnetoencephalography. Somatosensory change-related responses were evoked by an abrupt increase in stimulus strength in a train of current-constant square wave pulses at 100 Hz to the left median nerve at the wrist. Furthermore, auditory change-related responses were elicited by an increase in sound pressure by 15 dB in a continuous sound composed of a train of 25-ms pure tones. Binaural stimulation was used in Experiment 1, while monaural stimulation was used in Experiment 2. For both somatosensory and auditory stimuli, the conditioning and test stimuli were identical, and inserted at 2400 and 3000 ms, respectively. The results showed clear suppression of the test response in the bilateral parisylvian region, but not in the postcentral gyrus of the contralateral hemisphere in the somatosensory system. Similarly, the test response in the bilateral supratemporal plane (N100m) was suppressed in the auditory system. Furthermore, there was a significant correlation between suppression of right N100m and right parisylvian activity, suggesting that similar mechanisms are involved in both. Finally, a high test-retest reliability for suppression was seen with both modalities. Suppression revealed in the present study is considered to reflect sensory inhibition ability in individual subjects., Competing Interests: The authors have declared that no competing interests exist.

The opercular somatosensory region (OP) plays an indispensable role in pain perception. In the present study, we investigated the neurophysiological effects of transcranial direct current stimulation (tDCS) over the OP. Somatosensory-evoked magnetic fields following noxious intraepidermal electrical stimulation to the left index finger (pain-SEFs) were recorded before and after tDCS with a single-blind, sham-controlled, cross-over trial design. Three tDCS conditions of left anodal/right cathodal tDCS, left cathodal/right anodal tDCS (each, 2 mA, 12 min), and sham tDCS (2 mA, 15 s) were applied. Despite the subjective pain sensation being unaltered, the two anodal (real) interventions significantly decreased OP activity associated with pain-SEFs. In conclusion, tDCS over the OP with the present parameters did not have a significant impact on pain sensation, but modulated its cortical processing.

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.)

The sense of body ownership is based on integration of multimodal sensory information, including tactile sensation, proprioception, and vision. Distorted body ownership contributes to the development of chronic pain syndromes and possibly symptoms of psychiatric disease. However, the effects of disownership on cortical processing of somatosensory information are unknown. In the present study, we created a "disownership" condition in healthy individuals by manipulating the visual information indicating the location of the subject's own left hand using a mirror box and examined the influence of this disownership on cortical responses to electrical stimulation of the left index finger using magnetoencephalography (MEG). The event-related magnetic field in the right primary somatosensory cortex at approximately 50 msec (M50) after stimulus was enhanced under the disownership condition. The present results suggest that M50 reflects a cortical incongruence detection mechanism involving integration of sensory inputs from visual and proprioceptive systems. This signal may be valuable for future studies of the mechanisms underlying sense of body ownership and the role that disrupted sense of ownership has in neurological disease., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Objective: In contrast to tactile inputs, the organization and processing of nociceptive inputs in the primary somatosensory cortex (S1) remain largely unexplored. Few studies have examined the arrangement of nociceptive inputs in S1. The aim of this study was to investigate the representation of nociceptive inputs in the human cortex, including the somatosensory and posterior parietal cortices, from widely separated cutaneous sites., Methods: We examined the somatotopic organization of the nociceptive system in S1, opercular and posterior parietal cortices by measuring the magnetoencephalographic responses (somatosensory-evoked magnetic fields) of four healthy controls in response to intraepidermal electrical stimulation applied to the face, neck, back, elbow, wrist, hand, finger, knee, and foot, which selectively activated the Aδ fibers., Results: Magnetoencephalography demonstrated clear somatotopy in the S1 responses to noxious stimuli, with the foot representation in the extreme posteromedial position of S1 and the facial area in the extreme anterolateral position. There was little evidence of any clear somatotopic organization in the secondary somatosensory and posterior parietal cortices., Conclusion: These findings suggest that the nociceptive system uses the large body surface map in S1., Significance: This is the first MEG study to demonstrate the cortical representation of nociceptive inputs in the human S1. We showed that widely separated cutaneous sites clearly supported Penfield's homunculus., (Copyright © 2013 International Federation of Clinical Neurophysiology. Published by Elsevier Ireland Ltd. All rights reserved.)

Objective: We sought to clarify the presence of a temporal window of integration (TWI) in the somatosensory modality by manipulating the inter-stimulus interval (ISI)., Methods: We recorded cortical activity following the last of a train of electric pulses (stimulus offset) applied to the left hand in nine healthy volunteers using magnetoencephalography (MEG). Somatosensory evoked magnetic fields (SEFs) were elicited by the offset of a train of pulses 3s in total duration with four ISIs (25, 50, 75, and 100 ms)., Results: Results show that (i) off-M100 was clearly elicited by the ISI-25 and 50 ms conditions but not ISI-75 and 100 ms conditions, and (ii) the generator for off-M100 was mainly located in the contralateral primary and secondary somatosensory cortex (SI and SII)., Conclusion: The upper limit of the TWI in the somatosensory modality is between 50 and 75 ms, and the contralateral SI and SII play an important role in integrating temporal information., Significance: The present study clarifies the presence of the TWI in the somatosensory modality., (Copyright © 2011 International Federation of Clinical Neurophysiology. Published by Elsevier Ireland Ltd. All rights reserved.)

The somatic sensation of the tongue is necessary for daily life, but it is difficult to know the underlying neural mechanisms. In particular, because of the vomiting reflex and several morphological problems, no neuroimaging studies have examined somatosensory processing by stimulating the posterior part of the tongue, except for two magnetoencephalographic studies (Sakamoto et al., 2008a,b). This is the first study to clarify the human cortical processing for sensory perception by the posterior part of the tongue with a newly developed device and functional magnetic resonance imaging (fMRI). Stimulation of the left and right postero-lateral parts of the tongue induced significant activity in the primary somatosensory cortex (SI) and Brodmann area 40 in the right hemisphere and the anterior cingulate cortex (ACC). In contrast, antero-lateral stimulation produced activity only in the right SI. The activated region in SI was significantly larger following stimulation of the posterior than anterior part. These results indicate that a clear difference exists in somatosensory processing between stimulation of the antero-lateral and postero-lateral parts of the tongue, and a right hemisphere is dominant for the stimulation of both antero-lateral and postero-lateral areas. The activity in BA 40 and ACC may imply that the posterior of the tongue belongs to the visceral system.

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.

Quick detection of changes in the sensory environment is very important for survival, resulting in automatic shifts of attention to the event and the facilitation of subsequent processes to execute appropriate behaviors. The abrupt onset or offset of a sensory stimulus should also activate the neural network detecting changes. To test this hypothesis, we compared cortical on- and off-responses using somatosensory-evoked potentials (SEPs) elicited by a train of electrical pulses delivered to the right hand in eight healthy volunteers. SEPs were recorded from 15 electrodes on the scalp at three different interstimulus intervals (ISIs, 50, 20, and 10 ms) under two sets of conditions (attended and unattended). Both the onset and offset of stimulation evoked two similar components, P100 and N140, in the attended and unattended conditions. The latency of P100 and N140 in response to stimulus onset did not differ among the three ISIs, while the latency of both components in response to stimulus offset was significantly longer for the longer ISI; that is, detection of the cessation of the stimulation was based on short-term memory of the stimulus frequency. The present results supported a cortical network triggered by both the onset and offset of sensory stimulation. In this network, the change is automatically detected using a memory trace by comparing the abrupt event (on or off) with the preceding condition (silent or repetitive stimuli).

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.

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.

The go/nogo task is a useful paradigm for recording event-related potentials (ERPs) to investigate the neural mechanisms of response inhibition. In nogo trials, a negative deflection at around 140-300 ms (N2), which has been called the 'nogo potential', is elicited at the frontocentral electrodes, compared with ERPs recorded in go trials. In the present study, we investigated the generators of nogo potentials by recording ERPs and by using magnetoencephalography (MEG) simultaneously during somatosensory go/nogo tasks to elucidate the regions involved in generating nogo potentials. ERP data revealed that the amplitude of the nogo-N140 component, which peaked at about 155 ms from frontocentral electrodes, was significantly more negative than that of go-N140. MEG data revealed that a long-latency response peaking at approximately 160 ms, termed nogo-M140 and corresponding to nogo-N140, was recorded in only nogo trials. The equivalent current dipole of nogo-M140 was estimated to lie around the posterior part of the inferior frontal sulci in the prefrontal cortex. These results revealed that both nogo-N140 and nogo-M140 evoked by somatosensory go/nogo tasks were related to the neural activity generated from the prefrontal cortex. Our findings combining MEG and ERPs clarified the spatial and temporal processing related to somato-motor inhibition caused in the posterior part of the inferior frontal sulci in the prefrontal cortex in humans.

To elucidate the mechanisms underlying sensorimotor integration, we investigated modulation in the primary (SI) and secondary (SII) somatosensory cortices during the preparatory period of a self-initiated finger extension. Electrical stimulation of the right median nerve was applied continuously, while the subjects performed a self-initiated finger extension and were instructed not to pay attention to the stimulation. The preparatory period was divided into five sub-periods from the onset of the electromyogram to 3000 ms before movement and the magnetoencephalogram signals following stimulation in each sub-period were averaged. Multiple source analysis indicated that the equivalent current dipoles (ECDs) were located in SI and bilateral SII. Although the ECD moment for N 20 m (the upward deflection peaking at around 20 ms) was not significantly changed, that for P 30 m (the downward deflection peaking at around 30 m) was significantly smaller in the 0- to -500-ms sub-period than the -2000- to -3000-ms sub-period. As for SII, the ECD moment for the SII ipsilateral to movement showed no significant change, while that for the contralateral SII was significantly larger in the 0- to -500-ms sub-period than the -1500- to -2000-ms or -2000- to -3000-ms sub-period. The opposite effects of movement on SI and SII cortices indicated that these cortical areas play a different role in the function of the sensorimotor integration and are affected differently by the centrifugal process.

Objective: To investigate the somatotopic organization of the facial skin area in the secondary somatosensory cortex (SII) in humans., Methods: Somatosensory evoked magnetic fields following air-puff stimulation of 5 body sites, the foot, the lip and 3 points of the facial skin (forehead, cheek and mandibular angle point), were recorded. We focused on activities in SII following stimulation of these 5 sites and compared dipole locations among them., Results: There was a clear somatotopic organization in SII with lip in the most lateral area, foot in the most medial area and face in an intermediate area close to the lip area. However, there was no significant difference of dipole localization in SII among the 3 areas of facial skin, similar to the overlapped somatotopic organization of facial skin areas in the primary somatosensory cortex in our previous study., Conclusions: The facial skin areas are considered to occupy a small area in SII with insufficient spatial separation to differentiate each area of facial skin even using magnetoencephalography which has a high spatial resolution., Significance: This is the first systematic study of the activated regions in SII following stimulation of the facial skin.

The present study investigated the characteristics of the middle-latency negative potential of event-related potentials (ERPs) using somatosensory go/nogo tasks. We manipulated interstimulus interval (ISI) in Experiment 1 and stimulus probability in Experiment 2 and analyzed the subtracted difference waveform resulting from subtraction of the ERP evoked by the go stimulation from that evoked by the nogo stimulation. In Experiment 1, the peak latency of negativity became significantly longer as the ISI increased, but the peak amplitude was unchanged. The reaction time (RT) was longer with increasing ISI. In Experiment 2, manipulation of the stimulus probability yielded an increase in peak amplitude with decreasing probability of the nogo stimulus, but did not affect the latency. The RT increased as the probability of a nogo stimulus rose. Because manipulation of the ISI and stimulus probability elicited different brain activities, we hypothesized that manipulation of the ISI elicited a delay of the stimulus evaluation process including response inhibition, and that stimulus probability significantly affected the strength of the response inhibition process.

To investigate the representation of facial skin areas in the primary somatosensory cortex (SI), we recorded magnetic fields evoked by air pressure-induced tactile stimulation applied to six points on the face, lower lip and thumb. The thumb area in the SI was located more medial and superior to the lip area, which was consistent with Penfield's homunculus. However, the representations of all skin-covered areas including forehead, cheek, nose and chin in the SI were located between the thumb and lower lip area. There was no significant difference in location among the six facial points. Our results imply that lips occupy a large area of the face representation in the SI, whereas only a small area located between the thumb and lip areas is devoted to skin-covered surfaces. This is the first study showing that the facial skin areas in the human SI are located between the thumb and lower lip areas and close together.

Although numerous anatomical and electrophysiological findings in animal studies have supported a hierarchical scheme of somatosensory processing, precise activation timings of each cortical area are not known. Therefore we examined the temporal relationship of activities among multiple cortical areas using magnetoencephalography in humans. We found activations in Brodmann's areas 3b, 4, 1, 5 and the secondary somatosensory cortex region in the right hemisphere following transcutaneous electrical stimulation of the dorsum of the left hand. The mean onset latencies of each cortical activity were 14.4, 14.5, 18.0, 22.4 and 21.7 ms, respectively. The differences of onset latencies among these activations indicated the serial mode of processing both through the postcentral gyrus and through the primary and secondary somatosensory cortices.

Objective: We investigated the effects of a go/nogo task on event-related potentials (ERPs) evoked by somatosensory stimuli., Methods: ERPs following electrical stimulation of the second (go stimulus) or fifth (nogo stimulus) left-handed digit were recorded from 9 subjects. The recordings were conducted in 3 conditions: Control, Count and Movement. The subjects were instructed to count the go stimuli silently in Count, and respond to the go stimuli by grasping right hands in Movement. Go and nogo stimuli were presented at an even probability., Results: N140 was recorded in all conditions and P300 in Count and Movement. The mean amplitudes of the nogo stimuli in the interval 140-200 msec and nogo-N140 amplitude were significantly more negative than those of the go stimuli in Count or Movement. Nogo-P300 was larger in amplitude than go-P300 in Movement but not Count. The effect of P300 was applied to Fz and Cz, but not at Pz., Conclusions: In the present study, effects of a somatosensory go/nogo task on ERPs were investigated, and our findings were very similar to those of previous studies using visual and auditory go/nogo tasks. Therefore, we suggest that cortical activities relating to go/nogo tasks are not dependent on sensory modalities., Significance: The present study showed for the first time the go/nogo effects on somatosensory-evoked ERPs. These effects were similar to those in visual and auditory ERP studies.

To evaluate the effects of movement on cortical activities evoked by noxious stimulation, we recorded magnetoencephalography following noxious YAG laser stimulation applied to the dorsum of the left hand in normal volunteers. Results of the present study can be summarized as follows: (1) active movement of the hand ipsilateral to the side of noxious stimulation resulted in significant attenuation of both primary and secondary somatosensory cortices (SI and SII) in the hemisphere contralateral to the stimulated hand (cSI and cSII). Activity in the hemisphere ipsilateral to the side of stimulation (iSII) was not affected. (2) Active movement of the hand contralateral to the side of noxious stimulation resulted in significant attenuation of cSII. Activity in cSI and iSII was not affected. (3) Passive movement of the hand ipsilateral to the side of noxious stimulation resulted in significant attenuation of cSI. Activity in cSII and iSII was not affected. (4) Visual analogue scale (VAS) changes showed a similar pattern to the amplitude changes of cSII. These results suggest that activities in three regions are modulated by movements differently. Inhibition in cSI was considered to be mainly due to an interaction in SI by the signals ascending from the stimulated and movement hand. Inhibition in cSII was considered to be mainly due to particular brain activities relating to motor execution and/or movement execution associated with a specific attention effect. In addition, since VAS changes showed a similar relationship with the amplitude changes of cSII, cSII may play a role in pain perception.

We reported the changes of brain responses during sleep following auditory, visual, somatosensory and painful somatosensory stimulation by using magnetoencephalography (MEG). Surprisingly, very large changes were found under all conditions, although the changes in each were not the same. However, there are some common findings. Short-latency components, reflecting the primary cortical activities generated in the primary sensory cortex for each stimulus kind, show no significant change, or are slightly prolonged in latency and decreased in amplitude. These findings indicate that the neuronal activities in the primary sensory cortex are not affected or are only slightly inhibited during sleep. By contrast, middle- and long-latency components, probably reflecting secondary activities, are much affected during sleep. Since the dipole location is changed (auditory stimulation), unchanged (somatosensory stimulation) or vague (visual stimulation) between the state of being awake and asleep, different regions responsible for such changes of activity may be one explanation, although the activated regions are very close to each other. The enhancement of activities probably indicates two possibilities, an increase in the activity of excitatory systems during sleep, or a decrease in the activity of some inhibitory systems, which are active in the awake state. We have no evidence to support either, but we prefer the latter, since it is difficult to consider why neuronal activities would be increased during sleep.

To investigate the processing of noxious stimuli within the primary somatosensory cortex (SI), we recorded magnetoencephalography following noxious epidermal electrical stimulation (ES) and innocuous transcutaneous electrical stimulation (TS) applied to the dorsum of the left hand. TS activated two sources sequentially within SI: one in the posterior bank of the central sulcus and another in the crown of the postcentral gyrus, corresponding to Brodmann's areas 3b and 1, respectively. Activities from area 3b consisted of 20- and 30-ms responses. Activities from area 1 consisted of three components peaking at 26, 36 and 49 ms. ES activated one source within SI whose location and orientation were similar to those of the TS-activated area 1 source. Activities from this source consisted of three components peaking at 88, 98 and 109 ms, later by 60 ms than the corresponding TS responses. ES and TS subsequently activated a similar region in the upper bank of the sylvian fissure, corresponding to the secondary somatosensory cortex (SII). The onset latency of the SII activity following ES (109 ms) was later by 29 ms than that of the first SI response (80 ms). Likewise, the onset latency of SII activity following TS (52 ms) was later by 35 ms than that of area 1 of SI (17 ms). Therefore, our results showed that the processing of noxious and innocuous stimuli is similar with respect to the source locations and activation timings within SI and SII except that there were no detectable activations within area 3b following noxious stimulation.

We investigated the effects of sleep on pain-related somatosensory evoked magnetic fields (SEFs) following painful electrical stimulation to identify the mechanisms generating them in both fast A-beta fibers relating to touch and slow A-delta fibers relating to pain. While the subjects were awake, non-painful and painful electrical stimulations were applied, and while asleep, painful stimulation was applied to the left index finger. During awake, five components (1M-5M) were identified following both non-painful and painful stimulation, but the 4M and 5M at around 70-100 ms and 140-180 ms, respectively, were significantly enhanced following painful stimulation. During sleep, 1M and 2M generated in the primary somatosensory cortex (SI) did not show a significant change, 3M in SI showed a slight but significant amplitude reduction, and 4M and 5M generated in both SI and the secondary somatosensory cortex (SII) were significantly decreased in amplitude or disappeared. The 4M and 5M are complicated components generated in SI and SII ascending through both A-beta fibers and A-delta fibers. They are specifically enhanced by painful stimulation due to an increase of signals ascending through A-delta fibers, and are markedly decreased during sleep, because they much involve cognitive function.

Recent neuroimaging studies have shown that there are several brain areas that are activated by noxious stimuli, such as the primary and secondary somatosensory cortices. However, many of them are also activated by innocuous somatosensory stimuli, sensory stimuli other than the somatosensory system, or even tasks without sensory inputs, suggesting that the majority of brain activation in the so-called ‘pain matrix’ reflect event-related endogenous responses but not stimulus-related exogenous responses. For example, any somatosensory stimuli activate the primary and secondary somatosensory cortices, and any sensory stimuli activate the insula and the anterior cingulate gyrus. Therefore, pain-specific regions in the brain are not known. The physiological significance of well-known activations following noxious stimuli is an interesting matter, but to understand the mechanism of pain perception, the cortical nociceptive pathway starting with the primary nociceptive cortex and its origin in the thalamus should be determined at first. [ABSTRACT FROM AUTHOR]