stimuli restricted to elbow rotation; such neurons appear to be related to joint movement per se, independent of direction. 7. Differences between area 6 and area 4 cells related to elbow movements were barely significant. During active movements, average onset times of cells in areas 6 and 4 were essentially the same. A slightly larger proportion of area 6 cells responded to passive rotation of multiple joints, and during controlled elbow movements they more often exhibited complex response patterns. 8. These results, in conjunction with lesion and stimulation studies, are consistent with a sensory as well as motor role for precentral cortex neurons. Under passive conditions, the responses evoked by joint rotation and cutaneous stimulation may be utilized in perception of such stimuli, particularly in the absence of the more sensitive postcentral cells.
In awake monkeys we recorded activity of single "motor" cortex cells, four contralateral arm muscles, and elbow position, while operantly reinforcing several patterns of motor activity. With the monkey's arm held semiprone in a cast hinged at the elbow, we reinforced active elbow movements and tested cell responses to passive elbow movements. With the cast immobilized we reinforced isometric contraction of each of the four muscles in isolation, and bursts of cortical cell activity with and without simultaneous suppression of muscle activity. Correlations between a precentral cell and specific arm muscles consistently appeared under several behavioral conditions, but could be dissociated by reinforcing cell activity and muscle suppression.
Monkey motor cortex cells were recorded during isolated, isometric contractions of each of four representative arm muscles -- a flexor and extensor of wrist and elbow -- and comparable response averages computed. Most cells were coactivated with several of the muscles; some fired the same way with all four and others with none. Results suggest that many precentral cells have a higher order relation to muscles than motoneurons. Operantly reinforced bursts of cell activity were associated with coactivation of specific muscles, called the cell's "motor field"; the most strongly coactivated muscle was usually the one whose isolated contraction had evoked the most intense unit activity. During active elbow movements most cells fired in a manner consistent with their isometric patterns, but clear exceptions were noted. Differential reinforcement of unit activity and muscle suppression was invariably successful in dissociating correlations. The strength of each unit-muscle correlation was assessed by the relative intensity of their coactivation and its consistency under different response conditions. Several cells exhibited the most intense coactivation with the same muscle during all conditions. Thus, intensity and consistency criteria usually agreed, suggesting that strong correlations so determined may operationally define a "functional relation". However, correlations in the sense of covariation are neither necessary nor sufficient evidence to establish anatomical connections. To test the possibility of direct excitatory connections we stimulated the cortex, but found lowest threshold responses in distal muscles, even from points where most cells had been strongly correlated with proximal muscles. Post-spike averages of rectified EMG activity provided scant evidence for cell-related fluctuations in firing probabilities of any muscles.
The ability of human infants < or = 4 months of age to pursue objects smoothly with their eyes was assessed by presenting small target spots moving with hold-ramp-hold trajectories at ramp velocities of 4-32 deg/sec. Infants as young as 1 month old followed such target motions with a combination of smooth-pursuit and saccadic eye movements interrupted occasionally by periods when the eyes remained stationary. The slowest targets produced variable performance, but targets moving 8-32 deg/sec produced consistent pursuit behavior, even in the youngest infants. By the fourth month, eye-movement latency decreased and smooth-pursuit gain and the percentage of smooth pursuit per trial increased for all target velocities, though these measures had not yet reached adult levels.
The activity of jaw muscle receptors was studied by recording neurons in the mesencephalic nucleus of the trigeminal nerve in monkeys trained to control the position and movement of their mandible. Jaw position was measured by a weighted lever resting on the mandibular incisors. The force required to maintain the position of the lever was varied; in most cases it was either 25 or 360 g. Firing rates of neurons were related to stationary mandibular positions and to the velocity of movements during intervals when the movement velocity was constant. Of 49 neurons studied in detail, 21 fired at rates that were consistently and linearly related to static incisal openings. This static position sensitivity was typically about 5 spikes/mm of incisal opening. Most position-sensitive neurons fired at higher rates during opening movements and at lower rates during closing movements than would be accounted for by their position sensitivity. This sensitivity to the velocity of movement was not linear, however; slow closing movements sometimes did not produce a decrease in firing rate, and an actual increase during muscle shortening was seen in a few instances. The position sensitivity of eight neurons was evaluated during different loading conditions; in no case did it change substantially. Of the remaining 28 neurons, 26 fired at high rates during all opening movements and either stopped firing or fired at low, sporadic rates during closing movements. The static position sensitivity of these neurons was weak and variable both within and between neurons. The velocity sensitivity of these stretch-sensitive neurons was very nonlinear. Except for a range of slow movements (+/- 5 mm/s), the firing rate was maximal (200 spikes/s or higher) for most opening movements and zero for most closing movements. Maximal firing rates were higher when the loads being moved were increased from 25 to 360 g. The majority of position-sensitive neurons exhibited a large interspike-interval variability at wide incisal opening. In most of these neurons, this interspike-interval variability was periodic, usually at a rate of about 10 periods/s, and took the form of "saw-tooth" modulation on a record of instantaneous firing rate. Neurons that exhibited this modulation in a very prominent form also exhibited, in many instances, a substantial increase in firing rate during closing jaw movements.
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