Correlations between task-related activity and responses to perturbation in primate sensorimotor cortex.

1. Monkeys were trained to maintain hand position against a range of constant forces. Short-latency responses to passive wrist extension or flexion, as well as short-latency responses to stretch of a single wrist muscle, were recorded from units in areas 4, 3, 1, and 2. These responses were compared to unit activity during active holding and during active movement. 2. Units related to active holding and to active movement were most common in areas 4 and 2. Three-quarters of these units displayed a specific correlation between their passive and active behaviors. Thus, a unit excited by passive extension was excited during active holding against extension force and excited during an active flexion movement. This behavior is similar to the expected concurrent behavior of muscle stretch receptors. By demonstrating that a significant number of task-related units give qualitatively similar responses to passive extension and passive flexion, the results appear to explain the disagreement among previous studies (5, 9, 36) in regard to area 4 behavior during active and passive movements. 3. Area 4 units responded similarly to passive wrist extension and electromagnetic stretch of a single flexor muscle occurring in the absence of wrist extension, indicating that muscle stretch was important in determining area 4 unit responses to passive movements. 4. The similarity of area 4 behavior to area 2 behavior in active and passive situations, along with the observation that area 2 responses to passive movements occurred several milliseconds earlier than those of area 4, emphasizes the importance of area 2 in motor performance and is consistent with significant area 2 mediation of area 4 responses. 5. Results support the hypothesis of an oligosynaptic transcortical pathway (22, 32, 34), beginning in large part with muscle stretch receptors. Furthermore, the correlation noted between short-latency responses to passive movement and task-related activity suggests that this transcortical pathway not only mediates responses to passive movement but may be responsible, to a significant degree, for task-related activity during undisturbed performance. Thus, active position maintenance and active movement were probably accomplished, at least in part, by increasing and decreasing the influence of this pathway on specific area 4 neurons and thereby producing the patterns of area 4 activity responsible for task performance.