Centrifugal regulation of a task-relevant somatosensory signal triggering voluntary movement without a preceding warning signal

A warning signal followed by an imperative signal generates anticipatory and preparatory activities, which regulate sensory evoked neuronal activities through a top-down centrifugal mechanism. The present study investigated the centrifugal regulation of neuronal responses evoked by a task-relevant somatosensory signal, which triggers a voluntary movement without a warning signal. Eleven healthy adults participated in this study. Electrical stimulation was delivered to the right median nerve at a random interstimulus interval (1.75–2.25 s). The participants were instructed to extend the second digit of the right hand as fast as possible when the electrical stimulus was presented (ipsilateral reaction condition), or extend that of the left hand (contralateral reaction condition). They also executed repetitively extension of the right second digit at a rate of about 0.5 Hz, irrespective of electrical stimulation (movement condition), to count silently the number of stimuli (counting condition). In the control condition, they had no task to perform. The amplitude of short-latency somatosensory evoked potentials, the central P25, frontal N30, and parietal P30, was significantly reduced in both movement and ipsilateral reaction conditions compared to the control condition. The amplitude of long-latency P80 was significantly enhanced only in the ipsilateral reaction condition compared to the control, movement, contralateral reaction, and counting conditions. The long-latency N140 was significantly enhanced in both movement and ipsilateral reaction conditions compared to the control condition. In conclusion, short- and long-latency neuronal activities evoked by task-relevant somatosensory signals were regulated differently through a centrifugal mechanism even when the signal triggered a voluntary movement without a warning signal. The facilitation of activities at a latency of around 80 ms is associated with gain enhancement of the task-relevant signals from the body part involved in the action, whereas that at a latency of around 140 ms is associated with unspecific gain regulation generally induced by voluntary movement. These may be dissociated from the simple effect of directing attention to the stimulation.     

Mechanisms of differences in gating effects on short-and long-latency somatosensory evoked potentials relating to movement

We investigated the mechanisms underlying the differences in gating effects on short- and long-latency somatosensory evoked potentials (SEPs) relating to movement. SEPs were recorded in normal subjects for 6 different tasks in Experiment 1: Control, Movement, Distraction, Attention, Movement during Distraction and Movement during Attention, and for 4 different tasks in Experiment 2: Control, Passive Movement, Contralateral Movement and Movement Imagery. The amplitudes of short-latency SEPs were significantly reduced by active and passive movement of the stimulated hand, but long-latency SEPs (N140-P200) were significantly enhanced by active movement of the stimulated hand. Attention, Distraction, Contralateral Movement and Movement Imagery did not affect the amplitudes of SEPs. The degree of enhancement of long-latency SEPs by active Movement was greater than that by active movement with Attention or Distraction. Gating effects on long-latency SEPs were different from those on short-latency SEPs. Since this effect was not related to Attention/Distraction, Passive Movement, Movement Imagery or Movement of another site, it is probably due to specific centrifugal effects, which are different from more direct gating effects on short-latency components. This study showed the difference in gating effects on somatosensory perception depending on time periods following stimulation, which may indicate an interaction between motor and somatosensory cortex.     

Gating of somatosensory evoked potentials during voluntary movement of the lower limb in man

Somatosensory evoked potentials (SEPs) evoked by stimulation of the tibial nerve (TN) in the popliteal fossa, the sural nerve (Sur) at the lateral malleole, and an Achilles tendon (Achilles) tap were recorded before and during voluntary plantarflexion, dorsiflexion, and cocontraction of the ipsi- and contralateral foot in normal subjects. Suppression (gating) of the TN-SEP began around 60 ms before the onset of electromyographic activity (EMG), and became maximal 50–100 ms after the onset of EMG. Similar gating was observed for the SEP evoked by activation of muscle afferents (Achilles) and cutaneous afferents (Sur). The TN-SEP was similarly depressed at the onset of a plantarflexion as at the onset of dorsiflexion. A depression, although much smaller, was also observed at the onset of movement of the contralateral limb. The depression of the TN-SEP after the onset of EMG decreased when fast-conducting afferents were blocked by ischemia below the knee joint. The TN-SEP was equally depressed during tonic dorsiflexion, plantarflexion, and cocontraction of dorsi- and plantarflexors. The TN-SEP was depressed for up to 300 ms when preceded by stimulation of Sur or a biceps femoris tendon tap. Gating of lower limb SEPs thus appears to have both central and peripheral components of which neither seems to be specific for the muscle being contracted or the sensory afferents being stimulated. We encourage that caution is taken when drawing functional conclusions regarding movement-specific modulation of afferent inflow to the somatosensory cortex based on observations of gating of lower limb SEP. Received: 25 March 1997 / Accepted: 20 October 1997     

Facilitation of somatosensory evoked potentials by exploratory finger movements

Modification of somatosensory processing depending on the behavioral setting was studied. Active alternating movements of the fingers, passive tactile stimuli to the hand, and active exploration of objects were performed during recording of somatosensory evoked potentials (SEPs). SEPs were elicited by compound electrical median nerve stimulation and electrical stimulation at detection threshold of cutaneous median nerve fascicles identified by microneurography. Electrical stimulation was not time-locked to the studied condition.In comparison with SEPs at rest there was attenuation of early cortical potentials up to 25 ms post-trigger in all nonresting conditions. In stimulation of the compound median nerve as well as of isolated cutaneous fascicles of a hand actively exploring an object there was an additional increased negativity, peaking at 28 ms. This facilitory effect was independent of attentional focusing and was absent during exploration using the ipsilateral, non-electrically stimulated hand. In patients with parietal lesions the facilitatory effect was diminished on the affected side. Spline interpolated brain maps at this latency based on 32channel recordings in healthy volunteers showed a shift of local contralateral positive maximum from frontal to parietal during exploration, indicating enhancement of a tangential dipole. It is suggested that in conditions involving close sensorimotor interaction such as exploratory hand movements there is preactivation of a cortical area which is located in the central sulcus and receives cutaneous somatosensory inputs.     

Cortical representation of whole-body movement is modulated by proprioceptive discharge in humans

Previous studies have revealed the influence of ongoing sensory discharge on modulating the central representation of muscle afferents from individual limbs. In the present study, we explored the potential for such modulatory influence on the afferent discharge arising from induced whole-body movement. Vestibular and somato-sensory inputs arise from such whole-body movement. The convergence of these two modalities is important in motor control, especially for the maintenance of postural stability. We hypothesised that transmission of proprioceptive and vestibular information to the cortex would be reduced as a result of muscle-spindle discharge in knee extensor muscles. Perturbation-evoked responses (PERs), recorded from central scalp electrodes (C3, CZ, C4), were evoked through rapid translations of subjects who were seated in a chair on a movable platform. PERs were recorded during passive linear translations alone and preceded by vibration of the patellar tendon. The PER was characterised by a slow, negative potential peaking at approximately 150 ms (N150) following displacement of the chair. The amplitude of the PER was reduced following vibration to 56% of the control. Such reduction of PERs was comparable to the attenuation of somatosensory evoked potentials and soleus H-reflex magnitudes from tibial-nerve stimulation. We conclude that muscle-spindle discharge in knee extensor muscles leads to gating of both of these afferent pathways. These results have potential implications to the understanding of the CNS control of stability during ongoing movement.     

Stationarity of the somatosensory evoked potential

The validity of the somatosensory evoked potential for serial neurological evaluation requires an understanding of normal variations in the responses Studies were conducted in 15 normal adult volunteers during two nights of sleep and in five normal adult volunteers during waking hours. The response amplitudes and latencies obtained during the day were demonstrated to be stationary by use of the runs or sign test Similarly, the earliest portions of the scalp recorded evoked potential were stationary during sleep Later portions of the responses had significantly lower amplitudes and demonstrated increased latency during deep sleep. The stability of the early portions of the SSEP suggest that it is a reliable tool for serial neurological evaluation. In private practice     

Auditory and somatosensory event-related brain potentials in early blind humans

Previous event-related potential (ERP) studies have suggested a possible participation of the visual cortex of the blind in auditory processing. In the present study, somatosensory and auditory ERPs of blind and sighted subjects were recorded when subjects were instructed to attend to stimuli of one modality and to ignore those of the other. Both modalities were stimulated with frequent (standard) and infrequent (deviant) stimuli, which differed from one another in their spatial locus of origin. In the sighted, deviant stimuli of the attended modality elicited N2 type of deflections (auditory N2b and somatosensory N250) over the lateral scalp areas. In contrast, in the blind, these ERP components were centroposteriorly distributed, suggesting an involvement of posterior brain areas in auditory and somatosensory stimulus discrimination. In addition, the mismatch negativity, elicited by deviant auditory stimuli even when the somatosensory stimuli were attended, was larger in the blind than in the sighted. This appears to indicate enhanced automatic processing of auditory stimulus changes in the blind. Thus, the present data suggest several compensatory changes in both auditory and somatosensory modalities after the onset of early visual deprivation.     

脊髓体感与运动诱发电位术中联合监测的应用价值

目的：探讨脊髓体感诱发电位(SEP)与运动诱发电位(MEP)在脊髓手术中联合监测的临床应用价值。方法：对18例脊柱手术患者进行术中SEP和MEP联合监测，并用日本矫形学会量表(JOA)对患者术后神经功能进行评价。结果：全部患者术中SEP的P1、N1波幅有暂时性波动，潜伏期无明显变化。10例患者MEP的D1波波幅降低，但经改变手术方向后恢复正常，另8例患者MEP无明显变化。术后JOA评分较术前明显改善。结论：SEP及MEP术中联合监测，其波形稳定可靠，有利于避免“假阴性／假阳性”结果及术后神经功能障碍的发生。     

Interference of vibrations with input transmission in dorsal horn and cuneate nucleus in man: a study of somatosensory evoked potentials (SEPs) to electrical stimulation of median nerve and fingers

Summary The effects of 50 Hz palm vibrations on somatosensory potentials (SEPs) evoked by electrical stimulation of the median nerve at the wrist and of the 2nd and 3rd fingers were studied in 10 normal subjects. Vibrations were found to produce attenuation of the N13 spinal and P14 brainstem potentials and of the N20 contralateral parietal response. Brachial plexus (N9, P9) and dorsal column (P11) responses were not modified by vibrations. These SEP findings show: 1) that vibrations do not interfere at the periphery with the processing of brief ascending volleys triggered by an electrical stimulus and 2) that such an interference does occur in spinal dorsal horn and cuneate nucleus. Reduced input transmission in the cuneate nucleus is likely to be responsible for perceptual alterations induced by vibrations.     

New depth short-latency somatosensory evoked potential (SEP) component recorded in human SI area

To analyse short and long-latency (SEPs) recorded by chronically stereotactically electrodes implanted in SI area of two epileptic patients. Two drug-resistant epileptic patients (2 females, 38 and 15 years, respectively) suffering from left temporal and right frontal epilepsy respectively, were investigated by an electrode-chronically implanted in SI area. Short and long latency somatosensory evoked potentials were recorded by depth electrodes 10 days after implantation. This is the first study to describe a depth N36 response by an intracerebral recording electrode in the SI area, probably generated by a radially oriented generator, located in area 1. Furthermore, we confirmed a role of SI in the genesis of N60 component. Finally, our present data suggest that the SI area is still active at 120 ms after the stimulus, since in one patient (no. 2) we identified a N120 potential, reaching its maximal amplitude at the same depth as the N20 response.     

Modulation of somatosensory evoked responses in the primary somatosensory cortex produced by intracortical microstimulation of the motor cortex in the monkey

Summary Previous studies have shown that the amplitude of somatosensory evoked potentials is diminished prior to, and during, voluntary limb movement. The present study investigated the role of the motor cortex in mediating this movement-related modulation in three chronically prepared, awake monkeys by applying low intensity intracortical microstimulation (ICMS) to different sites within the area 4 representation of the arm. Air puff stimuli were applied to the contralateral arm or adjacent trunk at various delays following the ICMS. Somatosensory evoked potentials were recorded from the primary somatosensory cortex, areas 1 and 3b, with an intracortical microelectrode. The principal finding of this study was that very weak ICMS, itself producing at most a slight, localized, muscle twitch, produced a profound decrease in the magnitude of the short latency component of the somatosensory evoked potentials in the awake money. Higher intensities of ICMS (suprathreshold for eliciting electromyographic (EMG) activity in the target muscle, i.e. that muscle activated by area 4 stimulation) were more likely to decrease the evoked response and produced an even greater decrease. The modulation appeared to be, in part, central in origin since (i) it preceded the onset of EMG activity in 23% of experiments, (ii) direct stimulation of the muscle activated by ICMS, which mimicked the feedback associated with the small ICMS-induced twitch, was often ineffective and (iii) the modulation was observed in the absence of EMG activity. Peripheral feedback, however, may also make a contribution. The results also indicate that the efferent signals from the motor cortex can diminish responses in the somatosensory cortex evoked by cutaneous stimuli, in a manner related to the somatotopic order. The effects are organized so that the modulation is directed towards those neurones serving skin areas overlying, or distal to, the motor output.     

Primary somatosensory evoked magnetic fields elicited by sacral surface electrical stimulation

To explore the brain response to sacral surface therapeutic electrical stimulation (SSTES) for the treatment of refractory urinary incontinence and frequent micturition, evoked magnetic fields were measured in six healthy males. Electrical stimuli were applied between bilateral surface electrodes over the second through fourth posterior sacral foramens with intensity just below the pain threshold. Somatosensory evoked magnetic fields (SEFs) for the bilateral median (MN) and posterior tibial nerves (PTN) were also measured for the comparison. Sources of the early SEF peaks were superimposed on individual magnetic resonance images. The first peak latency for sacral stimuli, M30, occurred at 30.2 ± 0.8 ms (mean ± standard deviation, N = 6), with shorter latency than those for PTN stimulus (39.3 ± 1.4 ms, N = 12) and longer latency than those for MN stimulus (21.0 ± 0.9 ms, N = 12). The second peak latency for sacral stimuli, M50, occurred at 47.2 ± 2.9 ms (N = 6). Both M30 and M50 peaks showed a single dipole pattern over the vertex in the isofield maps. The equivalent current dipoles of M30 and M50 were both estimated near the medial end of the central sulcus with approximately posterior current direction. These results suggest that the sacral M30 and M50 are responses from the primary somatosensory cortex. The relatively long time lag between the onset and peak of M30 suggests that SSTES directly affects both the cauda equina and cutaneous nerve of the sacral surface.     

Cross‐modal relationships of neural gating with the subjective perception of respiratory and somatosensory sensations

Neural gating is a phenomenon whereby the response to a stimulus in the electroencephalogram (EEG) is attenuated when preceded by an identical stimulus. Attenuation of paired auditory clicks has repeatedly been shown to be affected in mental disorders, for example, schizophrenia. Neural gating has also been measured for respiratory and somatosensory sensations, however the attenuation of bodily relevant stimuli has not yet been systematically related to the subjective perception of bodily sensations. This research direction is potentially relevant to explaining disease trajectories in psychosomatic conditions characterized by chronic breathlessness and/or pain. In the present study, we recorded high‐density EEG from 85 healthy young adults while they experienced brief paired respiratory occlusions and brief paired electrocutaneous stimulation of the wrist. The event‐related potential N1 was measured centro‐laterally in response to the second relative to the first stimulus to quantify neural gating in both sensory domains. Participants experienced resistive loaded breaths and electrocutaneous stimuli of various intensities, rated their perceived intensity and unpleasantness, and performed magnitude estimation. Relationships of respiratory and somatosensory neural gating to the subjective intensity and unpleasantness of sensations, as well as the ability to discriminate sensations of varying intensities, were investigated intra‐modally and cross‐modally. We report significant relationships of the somatosensory neural gating to perceived intensity and unpleasantness of respiratory and somatosensory sensations, with the stronger neural gating relating to a stronger subjective intensity and unpleasantness. We discuss these unexpected findings through the lens of individual differences and different theoretical accounts on the origins of cortical attenuation of repetitive stimuli.     

Reproducibility and stability of neuroelectric source imaging in primary somatosensory cortex

The present study investigated the test-retest reliability of EEG source localization of somatosensory evoked potentials (SEPs) over an extended time period and tested the accuracy of source reconstruction co-registred with individual brain morphology (MRIs). Seven healthy subjects were stimulated pneumatically at the first digit and fifth digit of each hand and at the left and right lower corner of the mouth in two sessions spaced several weeks apart. At each location 1000 stimuli were presented. The overlay of the dipole localizations with the individual anatomic structure of the subjects' cortex was accomplished by the use of magnetic resonance images. A spherical 4-shell model of the head was used to localize the neuroelectric sources of the EEG data. In two cases a more realistically shaped 3 compartment model was computed using the boundary element method (BEM). The source localizations of the SEP component were found to be highly reproducible: the mean standard deviation of the dipole locations was 5.21 mm in the x-, 5.98 mm in the y- and 4.22 mm in the z-direction. BEM was not found to be superior to a 4-shell model. These data support the use of multi-electrode EEG recordings combined with MRI as an adequate method for the investigation of the functional organization of the somatosensory cortex.     

Reliability and validity of neuroelectric source imaging in primary somatosensory cortex of human upper limb amputees

The present study investigated the test-retest reliability of EEG source localizations of somatosensory evoked potentials (SEPs) in human upper limb amputees over a long time frame (several months) and examined the validity of source reconstruction. In two sessions spaced several months apart five unilateral upper limb amputees were stimulated at the first and fifth digit of the intact hand and at the left and right lower corner of the mouth. To examine the validity of the results of the neuroelectric source reconstruction a comparison with neuromagnetic source localization was performed for two subjects. The source localizations of the SEP components were found to be highly reproducible: the mean standard deviation of the dipole locations was 8.80 mm in the x-, 7.00 mm in the y- and 4.15 mm in the z-direction. The match of the comparison of EEG and MEG data was in the range of one centimeter. These results support the use of multi-electrode EEG recordings combined with MRI as an adequate method for the investigation of the functional organization of the somatosensory cortex in upper limb amputees and suggest high stability of cortical reorganization in these subjects.     

Changes in the centrifugal gating effect on somatosensory evoked potentials depending on the level of contractile force

In this study, we investigated the somatosensory evoked potentials (SEPs) during the preparatory period of self-initiated plantar flexion at different force levels of muscle contraction and elucidated the mechanism behind the centrifugal gating effect on somatosensory information processing. We recorded SEPs following stimulation of the tibial nerve at the popliteal fossa during the preparatory period of a 20% maximal voluntary contraction (MVC) and 50% MVC. The preparatory period was divided into two sub-periods based on the components of movement-related cortical potentials, the negative slope (NS sub-period) and the Bereitschaftspotential (BP sub-period). The subjects were instructed to concentrate on the movement and not to pay attention to the continuous electrical stimulation. Pre-movement SEPs were averaged separately during the two sub-periods under each MVC condition. The mean amplitudes of BP and NS were larger during the 50% MVC than the 20% MVC. As for the components of SEPs, during the NS sub-period the amplitude of P30 under the 50% MVC and N40 under both conditions were significantly smaller than that in the stationary sequence, and N40 amplitude was significantly smaller during the 50% MVC than the 20% MVC. During the BP sub-period, the amplitude of P30 and N40 during the 50% MVC was significantly smaller than during the stationary sequence, while it was not significantly different between the 20% and 50% MVCs. In conclusion, the extent of the centrifugal gating effect on SEPs was dependent on the activities of motor-related areas, which generated the NS and BP.     

Reduction of somatosensory evoked fields in the primary somatosensory cortex in a one-back task

In the present study, responses of the somatosensory cortex to sensory input of ten human volunteers were investigated during a one-back task with different conditions of attention. During an condition of attention subjects were requested to detect a predefined sequence of tactile stimuli applied to two different fingers of the dominant hand while a series of visual stimuli was presented simultaneously with an asynchronous stimulus-onset to the tactile stimuli. During an condition of distraction subjects received the identical series of visual and tactile stimuli like in the condition of attention but were now requested to detect a predefined stimulus sequence within the visual stimulus domain. In both conditions, somatosensory evoked magnetic fields (SEFs) to the tactile stimuli were recorded by means of a 31-channel magnetoencephalograph (MEG) from subjects‘ contralateral primary somatosensory cortex. The mean global field power, the dipole strength, the maximum current density, and the first component of the singular value decomposition (SVD) of magnetic fields were used to compare early components of the SEF in the conditions of attention versus distraction. Surprisingly, results revealed significant decreases of measures of all four parameters during the condition of attention as compared to the condition of distraction indicating that early responses of the primary somatosensory cortex became significantly reduced in the condition of attention. We hypothesize that changes in the centre-periphery-relationship of receptive fields in the primary somatosensory cortex may account for this unexpected result.     

Distinct effects of attention and affect on pain perception and somatosensory evoked potentials

The influence of affect and attention on sensory and affective pain as well as on somatosensory evoked potentials in response to painful and nonpainful electrical stimuli was investigated in a single experimental design. Affect was induced by pictures from the International Affective Picture System; attention was manipulated by asking participants to focus attention either on the pictures or on the electrical stimuli. Sensory and affective pain ratings were generally lower during exposure to positive compared to negative and neutral pictures. Attention modulated only sensory pain ratings with lower ratings with an attention focus on pictures than with an attention focus on sensory pain. The N150 was modulated by picture valence, the P260 by picture arousal. Furthermore, the P260 was modulated by attention with highest amplitudes with an attention focus on the stimulus intensity. This study provides neurophysiological evidence that attention and affect have distinct effects on pain processing.     

Movement-related potentials preceding voluntary movement are modulated by the mode of movement selection

In two experiments movement-related cortical potentials preceding voluntary movement were recorded. In experiment 1, subjects performed four motor tasks involving joystick movements. The four tasks differed in complexity (single vs sequential movements) and in the mode of movement selection, i.e., whether a movement or movement sequence was made in fixed or in self-determined directions. The choice of these tasks was based, firstly, on previous electrophysiological studies suggesting an effect of task-complexity on the amplitude of the readiness potential (RP) and, secondly, on previous positron emission tomography (PET) studies showing that activity of the supplementary motor area (SMA) is influenced by the mode of movement selection. The results show that, for single movements, RP amplitude is higher preceding freely selected movements than preceding movements in a fixed direction. In experiment 2 this effect was replicated using button presses instead of joystick movements. The results converge with PET evidence obtained in similar tasks and establish that the RP is modulated by the mode of movement selection. This modulation is probably related to differential involvement of the SMA.     

Localization of the human female breast in primary somatosensory cortex

Rationale: Despite an extensive body of research on the topography of the primary somatosensory cortex (S1) little is known about the representation of the trunk. Aim: The aim of this study was to determine the representation of the breast in S1 in human females. Results: The representation of the human breast in primary somatosensory cortex was determined in ten healthy female subjects. Non-painful electrical stimulation of the mammilla (Th4 dermatome), groin (L1 dermatome) and the first digit of both sides of the body activated cutaneous receptors and thus elicited somatosensory evoked potentials. The representation of these body parts in primary somatosensory cortex (S1) was determined using neuroelectric source imaging. Equivalent current dipole localizations were overlaid with individual structural magnetic resonance images to account for individual cortical differences. The breast representation was localized between the representation of the groin and the first digit. In the medial–lateral direction the representation of the breast was approximately 15 mm lateral of the longitudinal fissure in the contralateral hemisphere. Source localizations were stable across subjects. However, one subject showed ipsilateral representation of the breast, which might be related to bilateral receptive fields of the ventral body midline representation. This study confirms the Penfield and Rasmussen (1950) invasive data by use of noninvasive source imaging.