activity in primary motor and dorsal premotor cortex in reaching tasks with the contralateral versus ipsilateral arm. J Neurophysiol 89: 922-942, 2003; 10.1152/jn.00607.2002. To investigate the effector dependence of task-related neural activity in dorsal premotor (PMd) and primary motor cortex (M1), directional tuning functions were compared between instructed-delay reaching tasks performed separately with either the contralateral or the ipsilateral limb. During presentation of the instructional cue, the majority (55/90, 61%) of cells in PMd were tuned with both arms, and their dynamic range showed a trend for stronger discharge with the contralateral arm. Most strikingly, however, the preferred direction of most of these latter cells (41/55, 75%) was not significantly different between arms. During movement, many PMd cells continued to be tuned with both arms (53/90, 59%), with a trend for increasing directional differences between the arms over the course of the trial. In contrast, during presentation of the instructional cue only 5/74 (7%) cells in M1 were tuned with both arms. During movement, about half of M1 cells (41/74, 55%) were tuned with both arms but the preferred directions of their tuning functions were often very different and there was a strong bias toward greater discharge rates when the contralateral arm was used. Similar trends were observed for EMG activity. In conclusion, M1 is strongly activated during movements of the contralateral arm, but activity during ipsilateral arm movements is also common and usually different from that seen with the contralateral arm. In contrast, a major component of task-related activity in PMd represents movement in a more abstract or task-dependent and effector-independent manner, especially during the instructed-delay period. I N T R O D U C T I O NPerformance of reaching movements appears to require control at multiple levels of abstraction. For example, the neural mechanisms involved in deciding on the target for a reach need not necessarily take into account all the details of muscular contraction which must ultimately be controlled to accomplish the selected movement. Conversely, mechanisms involved in overt muscular control need not be sensitive to the criteria by which a particular action was selected. One therefore expects that different neural populations represent a given movement in different ways, emphasizing some cognitive, temporal, or spatial aspects while ignoring others.Indeed, many lines of evidence support a diversity of functions in reach-related areas of the cerebral cortex (for review, see Caminiti et al. 1998;Kalaska et al. 1997;Wise et al. 1997). For example, cells in primary motor cortex (M1) are strongly tuned to the direction, speed, and extent of movement (Caminiti et al. 1991;Crammond and Kalaska 1996;Crutcher and Alexander 1990;Fu et al. 1993;Georgopoulos 1991Georgopoulos , 1995Georgopoulos et al. 1982; Moran and Schwartz 1999) as well as to joint posture and muscle force Caminiti et al. 1991;Evarts 1968;Evarts et al. 1983;Kakei...
In instructed-delay (ID) tasks, instructional cues provide prior information about the nature of a movement to execute after a delay. Neuronal responses in dorsal premotor cortex (PMd) during the instructed-delay period (IDP) between the CUE and subsequent GO signals are presumed to reflect early planning stages initiated by the prior information. In contrast, in multiple-choice reaction-time (RT) tasks, all motor planning and execution processes must occur after the GO signal. These assumptions predict that neuronal planning correlates recorded during the IDP of ID trials should share common features with early post-GO activity in RT trials, and that those response components need not be recapitulated after the GO signal of ID trials. These two predictions were tested by comparing activity recorded in RT and ID tasks from 503 neurons in PMd and caudal (MIc) and rostral (MIr) primary motor cortex. The incidence and strength of directionally tuned IDP activity declined progressively from PMd to MIc. The directional tuning of activity during the IDP of ID trials was more similar to that in the reaction-time epoch (RTE) of RT trials than after movement onset, especially in PMd. A modulation of post-GO activity was often observed between RT and ID trials and was confined mainly to the RTE. This effect was also most prominent in PMd. The most common change was a reduction in intensity of short-latency phasic responses to the GO signal between RT and ID trials, especially in PMd cells with a short-latency phasic response to CUE signals. However, the largest group of cells in each area showed no large change in peak RTE activity between RT and ID trials, whether they were active in the IDP or not. Since early phasic CUE-related responses are least likely to be recapitulated after the GO signal in ID trials, they may be a neuronal correlate of an early planning stage such as response selection. Tonic IDP responses, which are not as strongly associated with a post-GO reduction in activity, may be related to other aspects of motor planning and preparation. Finally, a major component of the movement-related activity in both MI and PMd is not susceptible to modification by prior information and is indivisibly coupled temporally to movement execution.
Monkeys performed reaching movements in two opposite directions in a symmetrically rewarded GO/NOGO task with an instructed-delay period. Instructional cues were presented at the target locations. The decision not to move was clearly reflected in cell activity in dorsal premotor cortex, but not in parietal cortex area 5. In premotor cortex, the initial response (< 250 msec) of most cells to the appearance of the instructional cues in GO and NOGO trials was similar. However, by the end of the delay period, the responses of most cells were statistically different between the two trial types, and the population signals were much less directional in the NOGO trials than in the GO trials. In area 5, in contrast, single-cell and population signals were generally similar and strongly directional in both GO and NOGO trials. This result suggests a role for area 5 in visuomotor analysis for the guidance of limb movements. It further suggests that separate representations of potential motor responses to external inputs and of the intended response to that input can coexist in parietal and premotor cortex, respectively.
Because reaching movements have a clear objective--to bring the hand to the spatial location of an object--they are well suited to study how the central nervous system plans a purposeful act from sensory input to motor output. Most models of movement planning propose a serial hierarchy of analytic steps. However, the central nervous system is organized into densely interconnected populations of neurons. This paradox between the apparent serial order of central nervous system function and its complex internal organization is strikingly demonstrated by recent behavioral, modeling, and neurophysiological studies of reaching movements.
In this study human motor cortical activity was recorded with a customized micro-ECoG grid during individual finger movements. The quality of the recorded neural signals was characterized in the frequency domain from three different perspectives: (1) coherence between neural signals recorded from different electrodes, (2) modulation of neural signals by finger movement, and (3) accuracy of finger movement decoding. It was found that, for the high frequency band (60–120 Hz), coherence between neighboring micro-ECoG electrodes was 0.3. In addition, the high frequency band showed significant modulation by finger movement both temporally and spatially, and a classification accuracy of 73% (chance level: 20%) was achieved for individual finger movement using neural signals recorded from the micro-ECoG grid. These results suggest that the micro-ECoG grid presented here offers sufficient spatial and temporal resolution for the development of minimally-invasive brain-computer interface applications.
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