Corticomotoneuronal Research Articles

Direct and indirect corticospinal control of arm and hand motoneurons in the squirrel monkey (Saimiri sciureus).

Anatomic evidence suggests that direct corticomotoneuronal (CM) projections to hand motoneurons in the New World squirrel monkey (Saimiri sciureus) are weak or absent, but electrophysiological evidence is lacking. The nature of the corticospinal linkage to these motoneurons was therefore investigated first with the use of transcranial magnetic stimulation (TMS) of the motor cortex under ketamine sedation in five monkeys. TMS produced early responses in hand muscle electromyogram, but thresholds were high (compared with macaque monkey) and the onset latency was variable. Second, stimulation of the pyramidal tract (PT) was carried out with the use of chronically implanted electrodes in ketamine-sedated monkeys; this produced more robust responses that were markedly facilitated by repetitive stimulation, with little decrease in latency on the third compared with the first shock. Finally, postsynaptic potentials were recorded intracellularly from 93 arm and hand motoneurons in five monkeys under general chloralose anesthesia. After a single PT stimulus, the most common response was a small, slowly rising excitatory postsynaptic potential (EPSP), either alone (35 of 93 motoneurons) or followed by an inhibitory postsynaptic potential (39 of 93). The segmental delay of the early EPSPs was within the monosynaptic range (mean 0.85 ms); however, the rise time of these EPSPs was slow (mean 1.3 ms) and their amplitude was small (mean 0.74 mV). These values are significantly slower and smaller than EPSPs in a comparable sample of Old World macaque monkey motoneurons. The results show that CM connections do exist in the squirrel monkey but that they are weak and possibly located on the remote dendrites of the motoneurons. The findings are consistent with earlier anatomic studies. Repetitive PT stimulation produced large, late EPSPs in some motoneurons, suggesting that, in this species, there are relatively strong nonmonosynaptic pathways linking the corticospinal tract to hand motoneurons.

Effects on muscle activity from microstimuli applied to somatosensory and motor cortex during voluntary movement in the monkey.

It is well known that electrical stimulation of primary somatosensory cortex (SI) evokes movements that resemble those evoked from primary motor cortex. These findings have led to the concept that SI may possess motor capabilities paralleling those of motor cortex and speculation that SI could function as a robust relay mediating motor responses from central and peripheral inputs. The purpose of this study was to rigorously examine the motor output capabilities of SI areas with the use of the techniques of spike- and stimulus-triggered averaging of electromyographic (EMG) activity in awake monkeys. Unit recordings were obtained from primary motor cortex and SI areas 3a, 3b, 1, and 2 in three rhesus monkeys. Spike-triggered averaging was used to assess the output linkage between individual cells and motoneurons of the recorded muscles. Cells in motor cortex producing postspike facilitation (PSpF) in spike-triggered averages of rectified EMG activity were designated corticomotoneuronal (CM) cells. Motor output efficacy was also assessed by applying stimuli through the microelectrode and computing stimulus-triggered averages of rectified EMG activity. One hundred seventy-one sites in motor cortex and 68 sites in SI were characterized functionally and tested for motor output effects on muscle activity. The incidence, character, and magnitude of motor output effects from SI areas were in sharp contrast to effects from CM cell sites in primary motor cortex. Of 68 SI cells tested with spike-triggered averaging, only one area 3a cell produced significant PSpF in spike-triggered averages of EMG activity. In comparison, 20 of 171 (12%) motor cortex cells tested produced significant postspike effects. Single-pulse intracortical microstimulation produced effects at all CM cell sites in motor cortex but at only 14% of SI sites. The large fraction of SI effects that was inhibitory represented yet another marked difference between CM cell sites in motor cortex and SI sites (25% vs 93%). The fact that motor output effects from SI were frequently absent or very weak and predominantly inhibitory emphasizes the differing motor capabilities of SI compared with primary motor cortex.

Corticomotoneuronal contribution to the fractionation of muscle activity during precision grip in the monkey.

1. During independent finger movements, the intrinsic muscles of the hand show a fractionated pattern of activity in which the timing and amplitude of electromyographic (EMG) activity varies considerably from one muscle to another. It has been suggested that, in the macaque monkey, corticomotoneuronal (CM) cells that produce postspike facilitation (PSF) of EMG in these muscles contribute to this fractionation. To test this hypothesis, we have investigated the relationship between the pattern of PSF exerted by a CM cell and the pattern of activity shown by the cell and by its target muscles. 2. The activity of 15 identified CM cells was recorded from two monkeys that performed a precision grip task. Spike-triggered averaging of rectified EMG during the hold period of this task showed that each cell produced PSF in at least two intrinsic hand muscles. 3. Segments of data were selected from the initial movement period of the task in which the EMG activity in one target muscle was substantially greater than that of the other, and the mean firing rate of each CM cell was determined for these periods. 4. CM cells showed bursts of activity in the movement period. Most of them (13/15) had a significantly (P < 0.001) higher firing rate when one of its target muscles was more active than the other. For nine of these cells (identified as set A), this muscle was the one receiving the larger PSF. In four cases (set B), the reverse was true. Two cells (set C), which produced PSF of equal size in their target muscles, showed no change in firing rate across the periods of fractionated EMG activity. 5. All set A and set B cells fired at significantly (P < 0.001) higher rates during the movement period, in association with fractionation of EMG activity, than in the hold period, in which a cocontracted pattern of muscle activity was observed. 6. There were pronounced differences in the strength of PSF exerted by the CM cells on their target muscles during the fractionation periods. One CM cell exerted PSF of a muscle during one period of fractionation, but postspike suppression of the same muscle during the other period. 7. It is suggested that changes in the firing rate of a CM cell and in the degree of facilitation it exerts could both contribute to the fractionation of activity in its target muscles. Cells of set A appear to be specifically recruited in a manner that directly reflects the pattern of facilitation they exert on the sampled target muscles. These results may explain why the CM system is so important for the performance of relatively independent finger movements.

Primate rubromotoneuronal cells: parametric relations and contribution to wrist movement.

1. Fifty-nine rubromotoneuronal (RM) cells were identified in two rhesus monkeys on the basis of their postspike facilitation (PSpF) of rectified electromyographic (EMG) activity. These cells were studied in relation to a step tracking task requiring wrist movements between fixed target zones in flexion and extension. Movement away from a 0 position was opposed by spring-like loads (auxotonic). Additionally, nine cells were evaluated using an isometric task. Neuronal discharge could be divided into three basic components: background discharge in the absence of movement, phasic modulation during movement, and tonic modulation during sustained holding against external loads. 2. Four basic patterns of RM cell activity were observed in relation to ramp-and-hold wrist movements: phasic-tonic (44%), pure phasic (22%), pure tonic (2%), and unmodulated (24%). The discharge of unmodulated cells did not covary with movement parameters but, as with other RM cells, background discharge did increase in association with task performance. 3. The phasic discharge of RM cells led to the onset of target muscle EMG activity by an average of 89 +/- 82 ms (mean +/- SD, n = 104) in extensors and 88 +/- 74 ms (n = 30) in flexors. Target muscles are defined as ones showing PSpF of EMG activity. It was found that 94% of extensor and 87% of flexor RM cells discharged before or synchronous with the onset of target muscle EMG activity. 4. Thirty-one RM cells (53%) showed a tonic increase in cell activity during the static hold phase of the task. Twenty-three of these were tested for relations to static torque. Fifteen extension related cells and one flexion cell had significant, positive regression slopes for the relation between tonic discharge rate and static torque. The mean rate-torque slope for extension related cells was 160 Hz/Nm and 103 Hz/Nm for flexion related cells. These mean slopes are about one-third those of corticomotoneuronal (CM) cells. 5. Cell discharge rate was correlated with velocity and rate of change of torque (dT/dt) for 32 RM cells with a phasic component of discharge during movement. The peak increase in phasic discharge above tonic firing rate (PDI, peak dynamic index) was significantly correlated only with velocity in eight cells and only with dT/dt in five cells. The phasic discharge of four additional cells was correlated with both velocity and dT/dt, but for three of these cells, the correlation was stronger for velocity. The mean slope for the relation between velocity and PDI was 0.31 Hz.deg-1.s-1.(ABSTRACT TRUNCATED AT 400 WORDS)

Contribution of the monkey corticomotoneuronal system to the control of force in precision grip

1. The contribution of 33 corticomotoneuronal (CM) cells, recorded in the primary motor cortex, to the production of precision grip force has been investigated in four monkeys (Macaca nemestrina). These CM cells were shown, by spike-triggered averaging, to facilitate electromyographic (EMG) activity of hand and forearm muscles. 2. Single-cell recordings were obtained as the monkey performed a low force precision grip task under either isometric or auxotonic conditions. The monkey had to produce independent control of the forces exerted by the thumb and index finger and maintain them for 1-1.5 s. Steady force segments of data were selected trial-by-trial from these hold periods. For each segment the following mean values were determined: 1) CM cell firing rate, 2) EMG activity of facilitated muscles, and 3) index finger, thumb, and total force. 3. Of the 33 CM cells, 18 had a phasic-tonic pattern of discharge during the task, 7 were tonic, 5 had a ramplike increase, and 3 were deactivated during the hold period. 4. Of the 33 cells analyzed, 11 showed a significant positive (P < 0.05) correlation of their mean firing rate with static force; 4 of them had high correlation coefficients (P < 0.001). There was a considerable trial-by-trial variability in the cells' activity-force relationship. Six CM cells had significant negative correlations between their activity and isometric force (5 at the P < 0.001 level), showing lower firing rates with higher forces. 5. The force sensitivity of the CM cells, calculated from the rate-force slopes, was higher for either the thumb or the index finger force. Under isometric conditions the mean rate-force slopes, calculated from the best correlated digit force, was 32.4 Hz/N for eight positively correlated cells and -21.3 Hz/N for the cells with a negative correlation. 6. Correlation between CM cell spike activity and force was more common among neurons with slowly conducting axons (4/6 correlated) than for those with fast axons (13/27). 7. Significant correlations between target muscle EMG and force were always positive. The correlations between CM cell firing rate and target muscle EMG were comparable with those found between firing rate and force. Three of the CM cells with a negative correlation to force also had a negative correlation with EMG in one of their target muscles. 8. Each CM cell facilitated the EMG activity of one to five target muscles; postspike facilitation (PSF) was most common among intrinsic hand muscles (68/82 CM cell/muscle combinations).(ABSTRACT TRUNCATED AT 400 WORDS)

Response patterns and postspike effects of peripheral afferents in dorsal root ganglia of behaving monkeys.

1. The activity of single afferent units was recorded in cervical dorsal root ganglia (DRG) in two macaque monkeys as they generated alternating flexion and extension torques about the wrist during a step-tracking task. During these isometric and auxotonic muscle contractions, electromyographic (EMG) activity was recorded with electrode pairs in up to 12 independent forearm muscles. Spike-triggered averages (STAs) of rectified EMG activity were used to identify afferents that were associated with correlated facilitation of active muscles. 2. Our aim was to find peripheral afferents producing postspike effects in muscles and to compare their properties with those of corticomotoneuronal (CM) and rubromotoneuronal (RM) cells previously obtained under identical behavioral conditions. We documented the timing, magnitude and distribution of their postspike facilitation (PSF) of forearm muscles and investigated the response properties of task-related units. 3. Of 125 afferent units tested with STAs, 68 showed PSF of EMG activity in at least one muscle. Fifty-nine DRG units provided sufficiently long recordings to generate averages with greater than or equal to 2,000 triggers, the minimum number considered to demonstrate reliable effects. Of these 59 units, 29 (49%) were associated with facilitation of forearm muscle activity. 4. Many STAs showed a gradual increase in EMG activity starting before or near the afferent trigger spike; often superimposed on this broad facilitation was a sharply rising PSF starting at a longer latency. The earliest poststimulus facilitation evoked by single microstimuli delivered in DRG occurred in stimulus-triggered averages at a latency of 3.5 ms. In STAs the broad facilitation beginning at latencies shorter than the responses to electrical stimulation was attributed to synchronous discharges in other afferent units. The sharper postspike EMG increases occurring with latencies of greater than or equal to 3.5 ms were identified as PSF produced by the afferent. The PSF parameters documented in this study were measured after subtracting the effects of synchrony facilitation. 5. PSF of EMG activity began at a mean latency of 5.8 +/- 0.3 (SE) ms and peaked at a mean latency of 7.5 +/- 0.3 (SE) ms. In previous studies, the PSFs from CM and RM cells had mean onset latencies of 6.3 and 5.6 ms, respectively, and mean peak latencies of 10.2 and 9.1 ms. 6. A measure of the PSF amplitude is the mean percent increase (MPI), defined as the increase of the PSF above its base measured as a percentage of the prespike baseline mean.(ABSTRACT TRUNCATED AT 400 WORDS)

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)

Corticomotoneuronal connections in the rat: Evidence from double‐labeling of motoneurons and corticospinal axon arborizations

In order to investigate the possibility of direct corticomotoneuronal (CM) connections in the rat, an anterograde-retrograde double-labeling method was developed. Phaseolus vulgaris-leucoagglutinin (PHA-L) anterograde tracing of corticospinal axons was combined with retrograde labeling of spinal motoneurons either by a conjugate of choleragen subunit B with horseradish peroxidase (CB-HRP) or by wheat germ agglutinin (WGA). The location of PHA-L injection unilaterally in the forelimb area of sensorimotor cortex and the CB-HRP or WGA injections in corresponding contralateral wrist or digit extensors or flexors were determined and matched on the basis of movement responses elicited by intracortical microstimulation. Light microscopic observation showed, in addition to the main contralateral dorsal corticospinal tract (CST), the presence of four other CST minor components in the contralateral lateral, ipsilateral ventral, and ipsilateral dorsal funiculi of the cervical spinal white matter and at the base of contralateral dorsal horn of the gray matter, respectively. PHA-L-labeled CST axonal arbors were observed from Rexed's lamina I through lamina X of contralateral spinal gray matter, most extensively in laminae VI and VII; some CST axons reached the zone of motoneuronal somata in lamina IX and a few of them also entered the lateral and occasionally the ventral funiculi, ramifying in the white matter. Between the zones of PHA-L-labeled CST axonal arbors on the one hand and CB-HRP/WGA labeled spinal motoneuronal somata with their extensive dendritic trees on the other, there was a large overlap, covering partly both the gray and the white matter. PHA-L-labeled axonal boutons (en passant or terminaux) were seen to contact the dendrites or even the somata of motoneurons in the gray matter, according to light-microscopic criteria for identification of synaptic contacts. Axodendritic CM contacts were occasionally observed in the lateral funiculus of the white matter as well. In general, only a single contact was observed between an individual PHA-L-labeled CST axon and a given retrogradely labeled motoneuron. In contrast to the common notion that direct CM connections are a specialty of primates, the present morphological data support the presence of direct CM connections also in some other mammals, such as the rat.

Corticomotoneuronal postspike effects in averages of unrectified EMG activity.

1. Spike-triggered averaging (SpTA) of rectified electromyograms (EMGs) in awake monkeys is recognized as an effective means of establishing causal relations between cortical/cell discharge and muscle activity. The appearance of postspike facilitation (PSF) in averages of rectified EMG activity is interpreted as evidence of an underlying excitatory synaptic linkage between the trigger cell and motoneurons. Cells producing PSF are referred to as corticomotoneuronal (CM) cells. Similarly, postspike suppression (PSS) is interpreted as evidence of underlying inhibitory synaptic linkages. Studies to date have focused almost exclusively on averages of full-wave rectified EMG activity. Because the potential utility of SpTA of unrectified EMG activity has not been systematically examined, we compared postspike effects in SpTAs of rectified and unrectified EMGs for 44 cortical cells yielding 293 cell-target muscle pairs (CMPs). 2. Clear PSF was found in 110 of 293 averages of rectified EMG activity from 28 known CM cells. Forty-nine of these 110 CMPs (45%) also showed clear postspike effects (PSE) in the corresponding averages of unrectified EMGs activity. Loss of effects in averages of unrectified EMGs can be attributed to cancellation of the negative and positive phases of motor unit potentials. Mean onset latencies were similar for effects in rectified and unrectified EMGs [6.9 +/- 1.8 (SD) ms vs. 6.6 +/- 1.2 (SD) ms]. Overall, the magnitudes of effects measured as peak-to-noise ratios were also similar for effects in the two groups (8.8 vs. 8.7). The 61 CMPs that showed clear PSF in rectified EMGs but no effects in corresponding unrectified EMGs had, as a group, longer and more dispersed onset latencies and were weaker in magnitude than those that showed effects in both rectified and unrectified EMGs. Moreover, the occurrence of clear effects in averages of unrectified EMGs was correlated with the strength of PSF (75% of strong but only 29% of weak PSFs showed clear PSEs in averages of unrectified EMG activity). 3. Clear PSS was observed in 24 (8%) averages of rectified EMGs derived from nine cells. Remarkably, clear PSEs were found in 12 of the corresponding averages of unrectified EMG activity.(ABSTRACT TRUNCATED AT 400 WORDS)

Effects of synchrony between primate corticomotoneuronal cells on post-spike facilitation of muscles and motor units.

Cross-correlating the activity of neighboring motor cortex neurons recorded with independent electrodes in behaving monkeys has revealed synchronization peaks, largely due to common synaptic input. Corticomotoneuronal (CM) cells produced post-spike facilitation (PSF) of rectified forearm electromyograms (EMG); 15 cells synchronized with CM cells showed no PSF. Five pairs of CM cells with overlapping muscle fields exhibited similar synchrony peaks. The contribution of this synchrony to facilitation of common target muscles was assessed by two new methods: selective spike-triggered averaging and convolution. They showed that the PSF is composed predominantly of effects mediated by output of the triggering cell, but may include a broad, shallow component mediated by synchrony with other CM cells.

Encoding of motor parameters by corticomotoneuronal (CM) and rubromotoneuronal (RM) cells producing postspike facilitation of forelimb muscles in the behaving monkey

This paper compares the properties of corticomotoneuronal (CM) and rubromotoneuronal (RM) cells identified by postspike facilitation (PSF) of rectified EMG activity in the awake monkey. The postspike effects of CM and RM cells in flexors and extensors of the wrist and fingers have been determined, as have the discharge properties of these cells in relation to alternating ramp-and-hold wrist movements. The characteristics of postspike facilitation and postspike suppression (PSS) were similar for RM and CM cells. The magnitude of RM-PSF was weaker than CM-PSF and RM cells showed a stronger preference for facilitation of extensor muscles than CM cells. As with CM cells, the onset of discharge in RM cells preceded the onset of EMG activity in their target muscles. Tonic discharge related to static torque was more prominent in CM cells, whereas phasic discharge was more prominent in RM cells; however, many RM cells showed some tonic activity weakly related to static torque. We conclude that CM and RM cells share many common features; however, RM cells are concerned primarily with the dynamics of muscle contraction.

Activity of forelimb motor units and corticomotoneuronal cells during ramp-and-hold torque responses: comparisons with oculomotor cells

Publisher Summary To compare the functional organization of the somatomotor and oculomotor systems, it may be helpful to begin by comparing the response properties of their motoneurons and monosynaptically connected premotor neurons under behavioral conditions, which are as comparable as possible. In the somatomotor system, the activity of single forelimb motor units and input cells from precentral motor cortex, during ramp-and-hold torque responses has been sufficiently characterized to provide some basis for comparison. Such ramp-and-hold responses, alternating between flexion and extension target zones, were chosen to reveal the neural activity related to changes in force and to maintain static force; for purposes of discussion, these ramp-and-hold wrist responses may be considered analogous to saccadic eye movements followed by fixation. This chapter discusses the response properties and organization of somatic motor units and corticomotoneuronal (CM) cells, with brief comments on some interesting analogies with comparable oculomotor neurons. More information on the correlational linkages of the observed cells with their inputs and their targets would be helpful for understanding the source and consequences of their response properties.

Comparable patterns of muscle facilitation evoked by individual corticomotoneuronal (CM) cells and by single intracortical microstimuli in primates: evidence for functional groups of CM cells.

We compared the averaged responses of forelimb muscles to action potentials of single motor cortex cells and to single intracortical microstimuli (S-ICMS). Activity of precentral neurons and 12 identified forelimb muscles (6 flexors and 6 extensors of wrist and fingers) was recorded in macaques while they performed alternating ramp-and-hold wrist movements. Action potentials of cells that covaried reliably with wrist flexion or extension were used to compile spike-triggered averages (spike-TAs) of rectified electromyographic (EMG) activity of six synergistically coactivated muscles. Cells whose spikes were followed by a clear postspike facilitation (PSF) of rectified muscle activity were designated corticomotoneuronal (CM) cells. CM cells typically facilitated a subset of the coactivated muscles called the cell's target muscles. The relative strength of the PSF in different target muscles ranged from clear increases above base-line fluctuations to weak but significant effects. For each CM cell we characterized the "PSF profile" of facilitation across different muscles, defined as the relative strength of PSF in each of the coactivated agonist muscles. After identifying the CM cell's target muscles, we delivered S-ICMS through the microelectrode at the same site. Biphasic stimuli were delivered during the same wrist movements in which the recorded CM cell had been active. Stimulus intensities were too weak (typically 5-10 microA) and their repetition rate too slow (5-15 Hz) to evoke muscle excitation evident in the raw EMG record. However, stimulus-triggered averages (stimulus-TAs) of the rectified EMGs of coactivated muscles revealed consistent patterns of poststimulus facilitation (PStimF). In most cases the muscles facilitated by the CM cell in spike-TAs (n = 60) were also facilitated by S-ICMS in stimulus-TAs. At sites of CM cells the threshold stimulus intensities for evoking a statistically significant effect were between 0.5 and 2 microA. S-ICMS of 5 microA evoked PStimF that was, on the average, six times stronger than the PSF of the CM cell. The height of the facilitation peak relative to base-line fluctuations was 5-60 times greater for the stimuli than the spikes of the CM cell. The average onset latency of PStimF (8.0 +/- 1.2 ms) was 1.3 ms longer than the mean latency of PSF (6.7 +/- 1.4 ms). At two-thirds of the cortical sites where both spike- and stimulus-TAs were computed (n = 30), the PStimF profile exactly matched the PSF profile.(ABSTRACT TRUNCATED AT 400 WORDS)

Muscle Fields and Response Properties of Primate Corticomotoneuronal Cells

Publisher Summary This chapter summarizes the evidence concerning the distribution of postspike facilitation (PSF) in different forelimb muscles and the response properties of these cells during controlled ramp-and-hold wrist movements. Analysis of precentral cell responses during active and passive movement has revealed two major sources of input, central and peripheral; however, the functional significance of this activity remains uncertain as long as its output destinations are unknown. This chapter investigates those precentral cortex cells whose output effects on the activity of motor units could be confirmed by cross-correlation techniques. Spike-triggered averages (STA's) of rectified EMG activity have revealed that action potentials of certain precentral neurons are followed by a transient postspike facilitation (PSF) of average motor unit activity. In monkeys alternately flexing and extending the wrist against elastic loads, cortico-motoneuronal (CM) cells were identified by characteristic post-spike facilitation (PSF) of averaged forelimb muscle activity. The mean latency of such PSF was 6.7 ± 2.9 msec. The “muscle field” of such CM cells, defined as the set of muscles exhibiting PSF, could include only one muscle (30% of cells) or two (29%) or more (41%).

Noradrenaline in Acute Peripheral Circulatory Failure

How are the neurons that directly influence the motoneurons of a muscle distributed in the primary motor cortex (M1)? To answer this classical question we used retrograde transneuronal transport of rabies virus from single muscles of macaques. This enabled us to define cortico-motoneuronal (CM) cells that make monosynaptic connections with the motoneurons of the injected muscle. We examined the distribution of CM cells that project to motoneurons of three thumb and finger muscles. We found that the CM cells for these digit muscles are restricted to the caudal portion of M1, which is buried in the central sulcus. Within this region of M1, CM cells for one muscle display a remarkably widespread distribution and fill the entire mediolateral extent of the arm area. In fact, CM cells for digit muscles are found in regions of M1 that are known to contain the shoulder representation. The cortical territories occupied by CM cells for different muscles overlap extensively. Thus, we found no evidence for a focal representation of single muscles in M1. Instead, the overlap and intermingling among the different populations of CM cells may be the neural substrate to create a wide variety of muscle synergies. We found two additional surprising results. First, 15–16% of the CM cells originate from area 3a, a region of primary somatosensory cortex. Second, the size range of CM cells includes both “fast” and “slow” pyramidal tract neurons. These observations are likely to lead to dramatic changes in views about the function of the CM system.