Notes: Summary The discharge patterns of 144 single cortical neurones, within the cutaneous representation of the hand in area 2 (primary somatosensory cortex, SI), were studied in two rhesus monkeys during the performance of an active tactile discrimination task. These were compared to those previously described for units within areas 3b and 1 recorded from the same animals. The task consisted of making a single scanning movement of the digit tips over a surface (first half smooth; second half either smooth or rough). The nature of the texture encountered over the second half of the surface was indicated by the monkey making a differential lever response (push or pull) with the opposite hand. During the task, area 2 units with cutaneous receptive fields (RFs) on the digit tips of interest (those scanned over the surfaces) generally showed an increase in their discharge (75%); patterns of decreased discharge or no modulation (respectively, 12 and 13%) were rarely observed. Units with digital cutaneous RFs not in contact with the stimuli were much more likely to show either a pattern of decreased discharge or no modulation whatsoever (47% in each case), suggesting that there is some selection of cutaneous inputs in this task in that non-active inputs are selectively gated. For units with a cutaneous RF, the sign of modulation changed significantly across SI, in a manner consistent with a pattern of increased convergence onto the more caudal regions of SI. Overall, the proportions of area 2 units with digital RFs on the tips of interest that were classified as either texturerelated (25%) or movement-related (26%) were similar to those reported previously for areas 3b and 1, suggesting that their presumed roles in, respectively, the analysis of surface texture and the representation of the physical parameters of movement are shared and distributed across the three cytoarchitectonic subdivisions of SI under consideration. In addition, the discharge patterns of single texture-related cells in areas 3b, 1 and 2 did not reliably signal whether or not the animal successfully discriminated the surfaces, suggesting that information from a population of cells is required for the performance of the task. Texture-related responses in area 2 were, however, unique in two ways. Firstly, 35% of the texture-related units had additional discharges related to the performance of the scanning movement (texture- and movement-related cells); no such units were found in area 3b, and only one was encountered in area 1. Secondly, the texture-related responses of a subgroup of area 2 units (25%) varied as a function of the order of presentation of the surfaces. As response time also varied with the order of presentation, it is suggested that such cells might represent an intermediate step in the transformation of the sensory input to a behavioural response. Although most units with digital RFs were more responsive during active tactile discrimination than during passive movements of the digits over the same surfaces, secondary factors (speed of movement) were often responsible for this observation. In this aspect, area 2 discharge properties seemed closer to those previously described for area 3b than to those described for area 1. In contrast to both areas 3b and 1, however, the extent of modulation in the task of one-third of the units with larger multi-digit RFs was similar to that produced by classical RF testing, instead of being less (as found for areas 3b and 1). The latter observation suggests that some tactile inputs from the digit tips of interest are transmitted to area 2 relatively unchanged during exploratory movements, and that gating controls in this task of active tactile discrimination may be directed more towards cutaneous inputs to areas 3b and 1 than to area 2.

Notes: 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.

Notes: Summary The present experiments were designed to investigate the neuronal mechanisms, at the level of the primary somatosensory cortex, which underlie the observation that somatosensory cortical potentials evoked by air puff stimuli directed at the forearm are decreased, in a nonspecific and widespread manner, during voluntary movements about the elbow. Unitary discharge was recorded from 131 cells receiving cutaneous input from the hairy skin of the forearm or hand (areas 3b and 1) of two monkeys trained to perform rapid movements of the contralateral arm (elbow flexion or extension). Evoked unitary responses to air puff stimuli applied to the centre of the cell's receptive field, at various delays before and after the onset of movement, were recorded. Movement produced a significant decrease in the short latency excitatory response to the air puff in 89% of the cells (117/131); the remaining 11% were not modulated by movement. This movement-related “gating” of cutaneous inputs occurred regardless of the response pattern of the cells to movement alone, being observed in 91% of the cells with no movement-related discharge, and 89% of those with movement-related discharge. The air puff responses of cells with inputs from the forearm and the dorsum of the hand were all similarly modulated by movement and the modulation was clearly present prior to the onset of movement (mean onset, -66 ms). Variation in the depth of modulation as a function of the direction of the movement, flexion or extension, was observed in only a very small proportion of the modulated units (16/117); most showed no relationship to direction. It is suggested that, in this experimental situation, much of the modulation appears to occur at a pre-cortical level since there was no relationship between the pattern of discharge of cells in relation to movement alone and the pattern of movement-related gating of their responses to the air puff. Effects which might be consistent with a cortical origin for the modulation were only infrequently observed. The present results are strikingly similar to those obtained using the evoked potential method, and thus support the hypothesis that, in this task of rapid elbow movements, movement modulates the transmission of cutaneous signals from the hairy skin of the distal forelimb to primary somatosensory cortex in a nonspecific and widespread fashion.