Activity of spinal interneurons and their effects on forearm muscles during voluntary wrist movements in the monkey

We studied the activity of 577 neurons in the C-6-T-1 spinal cord of three awake macaque monkeys while they generated visually guided, isometric flexion/extension torques about the wrist. Spike-triggered averaging of electromyographic activity (EMG) identified the units' correlational linkages with less than or equal to 12 forearm muscles. One hundred interneurons produced changes in the level of average postspike EMG with onset latencies consistent with mono- or oligosynaptic connections to motoneurons; these were classified as premotor interneurons (PreM-INs). Most PreM-INs (82%) produced postspike facilitations in forearm muscles. Earlier spike-related features, often beginning before the trigger spike, were seen in spike-triggered averages from 72 neurons. Postspike effects were present in one muscle for 64% of the PreM-INs. Neurons with divergent linkages to larger "muscle fields" usually generated postspike effects in synergistic muscles. Fifty eight percent of the PreM-INs had postspike effects in flexor muscles only and 29% in extensor muscles only. Postspike effects were distributed relatively evenly among the primary flexor and extensor muscles studied. The mean percent change in EMG level from baseline and the mean onset latencies for postspike facilitations and postspike suppressions were similar. PreM-INs exhibited a variety of response patterns during the generation of isometric wrist torque. The response patterns and output effects of 24% of the PreM-INs were consistent with a strict reciprocal organization of flexor and extensor muscle control. For another 60% of the PreM-INs, there was a congruent relation between activity and output effects for only one direction of torque production. These neurons were active for both flexion and extension torques, including 37 neurons that exhibited bidirectional increases in discharge rate. The relatively small number of postspike suppressions observed suggests that inhibitory interneurons were silent when their target muscles were recruited. Compared with premotor neurons in the motor cortex, the red nucleus and the C-8-T-1 dorsal root ganglia, spinal PreM-INs affected flexor muscles in greater proportions and had smaller muscle fields. The magnitudes of postspike facilitations were similar in all premotor populations. Bidirectional activity, common for PreM-INs, was rare for corticomotoneuronal and premotor dorsal root ganglion cells, which discharge only for torques in their preferred direction.