Toshiaki Wasaka

Pain processing within the primary somatosensory cortex in humans

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 Brodmanns 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.

Centrifugal regulation of human cortical responses to a task-relevant somatosensory signal triggering voluntary movement

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.

Gating of somatosensory evoked magnetic fields during the preparatory period of self-initiated finger movement

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.

Somato-motor inhibitory processing in humans: a study with MEG and ERP.

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.

Movements modulate cortical activities evoked by noxious stimulation

&NA; 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.

Effects of a go/nogo task on event-related potentials following somatosensory stimulation

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.

Resource allocation and somatosensory P300 amplitude during dual task: effects of tracking speed and predictability of tracking direction

OBJECTIVE The amount of attentional resources allocated to a task is determined by the intrinsic demands, also denoted as task load or difficulty of the task. Effects of resource allocation on the somatosensory N140 and P300 were investigated in an inter-modal situation using a dual-task methodology. METHODS Under a dual-task condition, subjects concurrently performed a visuomotor tracking task and a somatosensory oddball task, while they performed just the oddball task under an oddball-only condition. In the tracking task, the subjects tracked the target line, which was presented on an oscilloscope and automatically moved, with the line which represented their own force generated by grip movement with the left hand. Tracking speed (experiment 1) and tracking predictability (experiment 2) were manipulated to vary task difficulty. N140, P300, and reaction time (RT) in the oddball task and tracking accuracy in the tracking task were measured. RESULTS The P300 and N140 amplitudes were reduced in the dual-task condition compared to those in the oddball-only condition. The fastest tracking speed produced lower tracking accuracy and later RT. However, the tracking speed did not affect the P300 or N140 amplitudes. In contrast, the P300 amplitude was smaller when the change in tracking direction was unpredictable than when it was predictable, without any differences in tracking accuracy or RT, N140. CONCLUSIONS The differences in behaviors among N140, P300, and RT following manipulation of task difficulty support the multiple-resource hypothesis, which defines functionally separate pools of resources. SIGNIFICANCE The present study may show that the P300 amplitude reflects modality-unspecific resource at more central level, and that the N140 amplitude involves perceptual resource.

Differential modulation in human primary and secondary somatosensory cortices during the preparatory period of self‐initiated finger movement

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 N20m (the upward deflection peaking at around 20 ms) was not significantly changed, that for P30m (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.

Higher anticipated force required a stronger inhibitory process in go/nogo tasks

OBJECTIVE We investigated the effect of the inhibitory process with increasing muscle force on event-related potentials (ERPs) and motor evoked potentials (MEPs). METHODS The subjects performed a S1-S2 paradigm with go/nogo tasks. S1 was an auditory tone burst, and S2 was an electrical stimulation applied to the second (go stimuli) or fifth digit (nogo stimuli) of the left hand. The recordings were conducted at 3 force levels; 10, 30 and 50% maximal voluntary contraction (MVC). After the presentation of S2, the subjects were instructed to adjust their force level to match the target line with a force trajectory line in only the go trials. RESULTS Nogo-N140 was significantly more negative in amplitude than go-N140 in all conditions, and became larger with increasing muscle force. The MEP, which was recorded at 150 ms after S2, became significantly smaller with increasing muscle force in nogo trials, whereas it became larger in go trials. CONCLUSIONS Our results indicated that stronger inhibitory cerebral activity was needed for a nogo stimulus, in the case where a stronger response was needed for a go stimulus. SIGNIFICANCE The present study showed a significant relationship between cortical inhibitory process and muscle force.

Effects of ISI and stimulus probability on event-related go/nogo potentials after somatosensory stimulation

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.