Bilateral force transients in the upper limbs evoked by single-pulse microstimulation in the pontomedullary reticular formation

Neurons in the pontomedullary reticular formation (PMRF) give rise to the reticulospinal tract. The motor output of the PMRF was investigated using stimulus-triggered averaging of electromyography (EMG) and force recordings in two monkeys ( M. fascicularis). EMG was recorded from 12 pairs of upper limb muscles, and forces were detected using two isometric force-sensitive handles. Of 150 stimulation sites, 105 (70.0%) produced significant force responses, and 139 (92.5%) produced significant EMG responses. Based on the average flexor EMG onset latency of 8.3 ms and average force onset latency of 15.9 ms poststimulation, an electromechanical delay of ∼7.6 ms was calculated. The magnitude of force responses (∼10 mN) was correlated with the average change in EMG activity ( P < 0.001). A multivariate linear regression analysis was used to estimate the contribution of each muscle to force generation, with flexors and extensors exhibiting antagonistic effects. A predominant force output pattern of ipsilateral flexion and contralateral extension was observed in response to PMRF stimulation, with 65.3% of significant ipsilateral force responses directed medially and posteriorly ( P < 0.001) and 78.6% of contralateral responses directed laterally and anteriorly ( P < 0.001). This novel approach permits direct measurement of force outputs evoked by central nervous system microstimulation. Despite the small magnitude of poststimulus EMG effects, low-intensity single-pulse microstimulation of the PMRF evoked detectable forces. The forces, showing the combined effect of all muscle activity in the arms, are consistent with reciprocal pattern of force outputs from the PMRF detectable with stimulus-triggered averaging of EMG.

Motor Outputs From the Primate Reticular Formation to Shoulder Muscles as Revealed by Stimulus-Triggered Averaging

The motor output of the medial pontomedullary reticular formation (mPMRF) was investigated using stimulus-triggered averaging (StimulusTA) of EMG responses from proximal arm and shoulder muscles in awake, behaving monkeys ( M. fascicularis). Muscles studied on the side ipsilateral (i) to stimulation were biceps (iBic), triceps (iTri), anterior deltoid (iADlt), posterior deltoid (iPDlt), and latissimus dorsi (iLat). The upper and middle trapezius were studied on the ipsilateral and contralateral (c) side (iUTr, cUTr, iMTr, cMTr). Of 133 sites tested, 97 (73%) produced a poststimulus effect (PStE) in one or more muscles; on average, 38% of the sampled muscles responded per effective site. For responses that were observed in the arm and shoulder, poststimulus facilitation (PStF) was prevalent for the flexors, iBic (8 responses, 100% PStF) and iADlt (13 responses, 77% PStF), and poststimulus suppression (PStS) was prevalent for the extensors, iTri (22 responses, 96% PStS) and iLat (16 responses, 81% PStS). For trapezius muscles, PStS of upper trapezius (iUTr, 49 responses, 73% PStS) and PStF of middle trapezius (iMTr, 22 responses, 64% PStF) were prevalent ipsilaterally, and PStS of middle trapezius (cMTr, 6 responses, 67% PStS) and PStF of upper trapezius (cUTr, 46 responses, 83% PStS) were prevalent contralaterally. Onset latencies were significantly earlier for PStF (7.0 ± 2.2 ms) than for PStS (8.6 ± 2.0 ms). At several sites, extremely strong PStF was evoked in iUTr, even though PStS was most common for this muscle. The anatomical antagonists iBic/iTri were affected reciprocally when both responded. The bilateral muscle pair iUTr/cUTr demonstrated various combinations of effects, but cUTr PStF with iUTr PStS was prevalent. Overall, the results are consistent with data from the cat and show that outputs from the mPMRF can facilitate or suppress activity in muscles involved in reaching; responses that would contribute to flexion of the ipsilateral arm were prevalent.

Plasticity in the Distribution of the Red Nucleus Output to Forearm Muscles After Unilateral Lesions of the Pyramidal Tract

It has been hypothesized that the magnocellular red nucleus (RNm) contributes to compensation for motor impairments associated with lesions of the pyramidal tract. To test this hypothesis, we used stimulus triggered averaging (StTA) of electromyographic (EMG) activity to characterize changes in motor output from the red nucleus after lesions of the pyramidal tract. Three monkeys were trained to perform a reach and prehension task. EMG activity was recorded from 11 forearm muscles including one elbow, five wrist, and five digit muscles. Microstimulation (20 μA at 20 Hz) was delivered throughout the movement task to compute StTAs. Two monkeys served as controls. In a third monkey, 65% of the left pyramidal tract had been destroyed by an electrolytic lesion method five years before recording. The results demonstrate a clear pattern of postlesion reorganization in red nucleus–mediated output effects on forearm muscles. The normally prominent extensor preference in excitatory output from the RNm (92% in extensors) was greatly diminished in the lesioned monkey (59%). Similarly, suppression effects, which are normally much more prominent in flexor than in extensor muscles (90% in flexors), were also more evenly distributed after recovery from pyramidal tract lesions. Because of the limited excitatory output from the RNm to flexor muscles that normally exists, loss of corticospinal output would leave control of flexors particularly weak. The changes in RNm organization reported in this study would help restore function to flexor muscles. These results support the hypothesis that the RNm is capable of reorganization that contributes to the recovery of forelimb motor function after pyramidal tract lesions.

Effects on wrist and digit muscle activity from microstimuli applied at the sites of rubromotoneuronal cells in primates

1. The purpose of this study was to use the techniques of spike- and stimulus-triggered averaging (SpTA and StTA, respectively) to examine the output organization of individual rubromotoneuronal (RM) cells in relation to clusters of neighboring cells. SpTA of electromyographic (EMG) activity in awake monkeys reveals the target muscles of an individual recorded neuron, whereas StTA reveals the target muscles of the neuronal aggregate activated by the stimulus. 2. Three questions were of particular interest. First, does the pattern of poststimulus facilitation (PStF) across forearm muscles match the pattern of postspike facilitation (PSpF)? Second, does the output of RM cell aggregates tested with StTA favor forearm extensor muscles, as reported for individual RM cells in the companion paper? Third, how do RM poststimulus effects compare with corticomotoneuronal (CM) poststimulus effects? 3. Microstimuli were applied at the sites of 37 RM cells, identified by SpTA of EMG activity in awake monkeys performing an alternating wrist movement task. 4. The magnitudes of PStF at 5, 10, and 20 microA were, respectively, 4.3, 10.1, and 13.7 times greater than PSpF of the same muscles, reflecting activation, by the stimulus, of multiple RM cells. RM cell PStF was weaker than CM PStF. 5. The onset latency of poststimulus suppression (PStS) exceeded that of PStF. For example, at 20 microA the difference was 2.6 ms, comparable with the difference between PSpF and postspike suppression (PSpS). 6. The patterns of poststimulus effects on forearm flexor and extensor muscles were categorized in the same manner as postspike effects. Three major patterns were observed: 1) pure facilitation, 2) reciprocal suppression, and 3) cofacilitation of extensors and flexors. 7. The profile of PStF across synergist muscles was broadly similar to that of PSpF. At 83% of sites, the muscle with the greatest PSpF was also the muscle with the greatest PStF. At 30% of sites (11 of 37), the set of muscles with PStF (muscle field) exactly matched the set with PSpF. Overall, the level of discrepancies in SpTAs versus StTAs computed at the same RM cell sites was 27%. Most of these discrepancies could be attributed to muscles with the weakest effects. 8. The fact that the poststimulus muscle fields at many RM cell sites matched the postspike fields at the same sites, even though the poststimulus effects were greater in magnitude and were mediated by more cells, suggests clustering of RM cells with similar target motoneuron pools.(ABSTRACT TRUNCATED AT 400 WORDS)