Postspike Facilitation Research Articles

Background firing in the auditory midbrain of the frog

Statistical characteristics of background firing in the midbrain auditory units of grass frog (Rana t. temporaria) located in torus semicircular (TS) were investigated. Only about 5% of the cells demonstrated prominent spontaneous firing. For such units the following characteristics were obtained: the distribution of interpulse intervals, the autocorrelation functions (ACF) for the real firing process and for the process with shuffled intervals, the hazard function (HF) and the joint distribution of adjacent interpulse intervals. The burstiness of firing was also estimated. In the absolute majority of the cells, the background firing demonstrated considerable deviation from the renewal process. There was weak but significant positive correlation between adjacent interpulse intervals. The burstiness of firing in the midbrain auditory units was moderate but higher than reported for medullary auditory neurons. The value of burstiness did not decrease after interval shuffling. Along with the reduction in excitability (generalized refractoriness) in many neurons observed post-spike facilitation effects were observed. Comparing background activity in medullary and midbrain nucleus suggests that there is an increase in complexity of the information processing along the auditory pathway.

Motion control of thumb and index finger of an artificial hand for precision grip using asynchronous decoding of CM cell activity

Recent brain machine interface (BMI) applications in humans [1] have shown the particular benefits of decoding of neural signals and of their use in motion control of artificial arms and hands for paralyzed people. Despite of the spectacular advances in the last decade on the kinematic control of reach and grasp movements, the dynamic control of these movements together with the choice of powerful decoding algorithms continues to be a major problem to be resolved. This work reports the results on asynchronous decoding, obtained with the application of artificial neural networks (ANN) [2], to both thumb and index finger kinematics and to their EMG. Neural data (spike trains and EMG) were recorded in the monkey. This work aims at providing a complete BMI framework to reproduce precision grip like hand movements with our 2 finger artificial hand (see Figure.1.A and B). The database was composed of up to six simultaneously recorded CM cell activity, of up to nine EMGs from different forearm muscles, and of the two fingertip position recorded in two monkeys while they performed a precision grip task (see Figure.1.A for robot reproduction). The CM cell activities were used as inputs and thus to train the time-delayed multi-layer perceptron (TDMLP) associated to each recording session in order to estimate both EMG and fingertip position signals. Each training epoch was performed with five different sliding window lengths that also determine the length of the input vector (i.e. the number of spikes considered at each instant) belonging to the time interval [25 ms,..,400 ms]. We trained the networks following three different paradigms: 1) Training the ANNs with only one spike train (from one CM cell). 2) Training the networks with all simultaneously recorded spike trains. 3) Only for the EMG estimation: training the networks with identified and non-identified CM cell spike trains. The identity of a cell as a CM cell was defined by the presence of a post spike facilitation (PSF) obtained by spike triggered averaging of the EMG. We then analyzed statistically the effects of each parameter on the estimating performance of the ANNs. Finally, we used the fingertip position signals estimated by the corresponding trained ANN to drive the index (4 DoF) and thumb finger (5 DoF) of our artificial hand (Shadow Robot Company ©). We then compared the reproduction performance of the hand to the recorded and estimated position signals (Figure ​(Figure1B1B). Figure 1 A. Artificial forearm with a 2-finger hand of the precision grip setup actuated by pneumatic muscles, B. Reproduction of thumb and index finger position.

The activity of primary motor cortex corticospinal neurons during tool use by macaque monkeys.

It has been suggested that the distinctive capacity of some nonhuman primates to use tools may reflect a well-developed corticospinal system and, in particular, direct cortico-motoneuronal (CM) connections to hand muscles. We investigated the activity of corticospinal neurons in the primary motor cortex hand area during the use of a tool by two adult macaque monkeys. They used a light rake to retrieve food rewards placed in their extrapersonal space. An analysis of EMG activity showed that the rake task involved a complex interaction of muscles acting on the digits, hand, and arm. Sixty-nine corticospinal neurons were identified antidromically as pyramidal tract neurons (PTNs). When tested on the rake task, most (64 of 69; 93%) showed a significant modulation of their discharge during at least one of three task periods: grasping the rake, projecting it beyond the food reward, and then pulling it back to retrieve the reward. Discharge patterns were heterogeneous, and many PTNs showed significant suppression of discharge during raking. Seventeen of the 69 PTNs recorded during the rake task were further identified as CM cells, exerting clear postspike facilitation on digit muscles, demonstrating that the CM system contributes to the skilled use of tools. We compared the activity of each PTN on the rake task with that during precision grip. Most PTNs (90%) modulated their activity significantly for both tasks, demonstrating that PTNs activated by a task involving fractionated movements of the digits are also recruited during rake use, although there were often contrasting patterns of PTN recruitment and muscle activity for the two tasks.

A neural basis for motor primitives in the spinal cord.

Motor primitives and modularity may be important in biological movement control. However, their neural basis is not understood. To investigate this, we recorded 302 neurons, making multielectrode recordings in the spinal cord gray of spinalized frogs, at 400, 800, and 1200 mum depth, at the L2/L3 segment border. Simultaneous muscle activity recordings were used with independent components analysis to infer premotor drive patterns. Neurons were divided into groups based on motor pattern modulation and sensory responses, depth recorded, and behavior. The 187 motor pattern modulated neurons recorded comprised 14 cutaneous neurons and 28 proprioceptive neurons at 400 mum in the dorsal horn, 131 intermediate zone interneurons from approximately 800 microm depth without sensory responses, and 14 motoneuron-like neurons at approximately 1200 microm. We examined all such neurons during spinal behaviors. Mutual information measures showed that cutaneous neurons and intermediate zone neurons were related better to premotor drives than to individual muscle activity. In contrast, proprioceptive-related neurons and ventral horn neurons divided evenly. For 46 of the intermediate zone interneurons, we found significant postspike facilitation effects on muscle responses using spike-triggered averages representing short-latency postspike facilitations to multiple motor pools. Furthermore, these postspike facilitations matched significantly in both their patterns and strengths with the weighting parameters of individual primitives extracted statistically, although both were initially obtained without reference to one another. Our data show that sets of dedicated interneurons may organize individual spinal primitives. These may be a key to understanding motor development, motor learning, recovery after CNS injury, and evolution of motor behaviors.

Do Corticomotoneuronal Cells Predict Target Muscle EMG Activity?

Data from two rhesus macaques were used to investigate the pattern of cortical cell activation during reach-to-grasp movements in relation to the corresponding activation pattern of the cell's facilitated target muscles. The presence of postspike facilitation (PSpF) in spike-triggered averages (SpTAs) of electromyographic (EMG) activity was used to identify cortical neurons with excitatory synaptic linkages with motoneurons. EMG activity from 22 to 24 muscles of the forelimb was recorded together with the activity of M1 cortical neurons. The extent of covariation was characterized by 1) identifying the task segment containing the cell and target muscle activity peaks, 2) quantifying the timing and overlap between corticomotoneuronal (CM) cell and EMG peaks, and 3) applying Pearson correlation analysis to plots of CM cell firing rate versus EMG activity of the cell's facilitated muscles. At least one firing rate peak, for nearly all (95%) CM cells tested, matched a corresponding peak in the EMG activity of the cell's target muscles. Although some individual CM cells had very strong correlations with target muscles, overall, substantial disparities were common. We also investigated correlations for ensembles of CM cells sharing the same target muscle. The ensemble population activity of even a small number of CM cells influencing the same target muscle produced a relatively good match (r >/= 0.8) to target muscle EMG activity. Our results provide evidence in support of the notion that corticomotoneuronal output from primary motor cortex encodes movement in a framework of muscle-based parameters, specifically muscle-activation patterns as reflected in EMG activity.

Functional relations between primate motor cortex cells and muscles: fixed and flexible.

In behaving monkeys the effects of motor cortex cells on muscles are inferred from two quite different types of 'correlational' evidence: their coactivation and cross-correlation. Many precentral cells are coactivated with limb muscles, suggesting that they make a proportional contribution to muscle activity; however, such coactivation is typically quite flexible, and can be changed by operantly conditioning the dissociation of cell and muscle activity. Cross-correlating cells and muscles by spike-triggered averaging of the electromyogram (EMG) shows that certain cells produce short-latency post-spike facilitation of EMG; this correlational linkage is relatively fixed under different behavioural conditions and its time course suggests it is mediated by a corticomotoneuronal (CM) synaptic connection. CM cells typically facilitate a set of coactivated agonist muscles, and some also inhibit their antagonists. The firing patterns of CM cells can differ significantly from those of their target muscles. During ramp-and-hold wrist responses most CM cells discharge a phasic burst that precedes target muscle onset and that contributes to changes in muscle activity. At low force levels many CM cells are activated without their target motor units. Conversely, many CM cells are paradoxically inactive during rapid forceful movements that vigorously activate their target muscles; they appear to be preferentially active during finely controlled movements. Thus CM cells, with a fixed correlational linkage to their target muscles, may be recruited without their target muscles, and vice versa.

Descending Signals From the Pontomedullary Reticular Formation Are Bilateral, Asymmetric, and Gated During Reaching Movements in the Cat

We examined the contribution of neurons within the pontomedullary reticular formation (PMRF) to the control of reaching movements in the cat. We recorded the activity of 127 reticular neurons, including 56 reticulospinal neurons, during movements of each forelimb; 67/127 of these neurons discharged prior to the onset of activity in the prime flexor muscles during the reach of the ipsilateral limb and form the focus of this report. Most neurons (63/67) showed similar patterns and levels of discharge activity during reaches of either limb, although activity was slightly greater during reach of the ipsilateral limb. In 26/67 cells, the initial change in discharge activity was time-locked to the go signal during reaches of either limb; we have argued that this early discharge contributes to the anticipatory postural adjustments that precede movement. In 11/26 cells, the initial change in activity was reciprocal for reaches with the left and right limbs, although activity during the movement was nonreciprocal. Spike-triggered averaging produced postspike facilitation or depression (PSD) in 12/50 cells during reaches of the limb ipsilateral to the recording site and in 17/49 cells during reach of the contralateral limb. Some cells produced PSD in ipsilateral extensor muscles before the start of the reach and during reaches made with the contralateral, but not the ipsilateral limb; this suggests the signal must be differentially gated. Overall, the results suggest a strong bilateral, albeit asymmetric, contribution from the PMRF to the control of posture and movement during voluntary movement.

Afferent Encoding of Central Oscillations in the Monkey Arm

We have investigated whether peripheral afferent fibers could encode the central oscillations that are commonly seen in the primate motor system. We analyzed 52 single afferent recordings from the C8/T1 dorsal root ganglia of two monkeys performing an isometric wrist flexion-extension task. Coherence and directed coherence were calculated between the afferent spikes and forearm EMG. Seven of 52 cells were identified as Group Ia afferents by the production of narrow postspike facilitation in spike-triggered averages of rectified EMG. These identified afferents showed significant coherence, and directed coherence, with EMG over a wide frequency range. By contrast, coherence was weak for a population that showed little directional preference for flexion or extension movements during task performance, and probably contained mainly cutaneous afferents. Oscillations are known to appear in muscle activity; their presence in afferent firing as well implies that central oscillations pass around a peripheral feedback loop and may be involved in sensorimotor integration.

Synchrony between Neurons with Similar Muscle Fields in Monkey Motor Cortex

Synchronous firing of motor cortex cells exhibiting postspike facilitation (PSF) or suppression (PSS) of hand muscle EMG was examined to investigate the relationship between synchrony and output connectivity. Recordings were made in macaque monkeys performing a precision grip task. Synchronization was assessed with cross-correlation histograms of the activity from 144 pairs of simultaneously recorded neurons, while spike-triggered averages of EMG defined the muscle field for each cell. Cell pairs with similar muscle fields showed greater synchronization than pairs with nonoverlapping fields. Furthermore, cells with opposing effects in the same muscles exhibited negative synchronization. We conclude that synchrony in motor cortex engages networks of neurons directly controlling the same muscle set, while inhibitory connections exist between neuronal populations with opposing output effects.

Correlations between corticomotoneuronal (CM) cell postspike effects and cell-target muscle covariation.

The presence of postspike facilitation (PSpF) in spike-triggered averages of electromyographic (EMG) activity provides a useful means of identifying cortical neurons with excitatory synaptic linkages to motoneurons. Similarly the presence of postspike suppression (PSpS) suggests the presence of underlying inhibitory synaptic linkages. The question we have addressed in this study concerns the extent to which the presence and strength of PSpF and PSpS from corticomotoneuronal (CM) cells correlates with the magnitude of covariation in activity of the CM cell and its target muscles. For this purpose, we have isolated cells during a reach and prehension task during which the activity of 24 individual proximal and distal forelimb muscles was recorded. These muscles show broad coactivation but with a highly fractionated and muscle specific fine structure of peaks and valleys. Covariation was assessed by computing long-term (2 s) cross-correlations between CM cells and forelimb muscles. The magnitude of cross-correlations was greater for muscles with facilitation effects than muscles lacking effects in spike-triggered averages. The results also demonstrate a significant relationship between the sign of the postspike effect (facilitation or suppression) and the presence of a peak or trough in the cross-correlation. Of all the target muscles with facilitation effects in spike-triggered averages (PSpF, PSpF with synchrony, or synchrony facilitation alone), 89.5% were associated with significant cross-correlation peaks, indicating positively covarying muscle and CM cell activity. Seven percent of facilitation effects were not associated with a significant effect in the cross-correlation, whereas only 3.4% of effects were associated with correlation troughs. In contrast, of all the muscles with suppression effects in spike-triggered averages, 38.9% were associated with significant troughs in the cross-correlation, indicating an inverse relation between CM cell and muscle activity consistent with the presence of suppression. Fifty-five percent of suppression effects was associated with correlation peaks, whereas 5.6% was not associated with a significant effect in the cross-correlation. Limiting the analysis to moderate and strong facilitation effects, the magnitude of PSpF was correlated weakly with the magnitude of the cell-muscle cross-correlation peak. Nevertheless, the results show that although many CM cell-target muscle pairs covary during the reach and prehension task in a way consistent with the sign and strength of the CM cell's synaptic effects on target motoneurons, many exceptions exist. The results are compatible with a model in which control of particular motoneuron pools reflects not only the summation of signals from many CM cells but also signals from additional descending, sensory afferent, and intrinsic spinal cord neurons. Any one neuron will make only a small contribution to the overall activity of the motoneuron pool. In view of this, it is not surprising that relationships between postspike effects and CM cell-target muscle covariation are relatively weak with many apparent incongruities.

Corticomotoneuronal postspike effects in shoulder, elbow, wrist, digit, and intrinsic hand muscles during a reach and prehension task.

We used spike-triggered averaging of rectified electromyographic activity to determine whether corticomotoneuronal (CM) cells produce postspike effects in muscles of both proximal and distal forelimb joints in monkeys performing a reach and prehension task. Two monkeys were trained to perform a self-paced task in which they reached forward from a starting position to retrieve a food reward from a small cylindrical well. We compiled spike-triggered averages from 22 to 24 separate forelimb muscles at both proximal (shoulder, elbow) and distal (wrist, digits, intrinsic hand) joints. Of 174 cells examined, 112 produced postspike effects in at least one of the target muscles. Of those cells, 45.5% produced postspike effects in both proximal and distal forelimb muscles. A nearly equal number (44.7%) produced postspike effects in distal muscles only, whereas a clear minority (9.8%) produced postspike effects in only proximal muscles. The majority of CM cells (71.4%) produced effects in two or more muscles, with an average muscle field of 3.1 +/- 2.1 (mean +/- SD) for facilitation plus suppression. Of 345 postspike effects identified, 70.7% were facilitation effects and 29.3% were suppression effects. The large majority of effects (72.2%) were in distal muscles. When averaged by joint, the latency and peak magnitude of postspike facilitation showed a stepwise increase from proximal to distal joints. The results of this study show that the majority of CM cells engaged in coordinated forelimb reaching movements facilitate and/or suppress muscles at multiple joints, including muscles at both proximal and distal joints. The results also show that CM cells make more frequent and more potent terminations in motoneuron pools of distal compared with proximal muscles.

Computer simulation of post-spike facilitation in spike-triggered averages of rectified EMG.

When the spikes of a motor cortical cell are used to compile a spike-triggered average (STA) of rectified electromyographic (EMG) activity, a post-spike facilitation (PSF) is sometimes seen. This is generally thought to be indicative of direct corticomotoneuronal (CM) connections. However, it has been claimed that a PSF could be caused by synchronization between CM and non-CM cells. This study investigates the generation of PSF using a computer model. A population of cortical cells was simulated, some of which made CM connections to a pool of 103 motoneurons. Motoneurons were simulated using a biophysically realistic model. A subpopulation of the cortical cells was synchronized together. After a motoneuron discharge, a motor unit action potential was generated; these were summed to produce an EMG output. Realistic values were used for the corticospinal and peripheral nerve conduction velocity distribution, for slowing of impulse conduction in CM terminal axons, and for the amount of cortical synchrony. STA of the rectified EMG from all cortical neurons showed PSF; however, these were qualitatively different for CM versus non-CM cells. Using an epoch analysis to determine reliability in a quantitative manner, it was shown that the onset latency of PSF did not distinguish the two classes of cells after 10,000 spikes because of high noise in the averages. The time of the PSF peak and the peak width at half-maximum (PWHM) could separate CM from synchrony effects. However, only PWHM was robust against changes in motor unit action-potential shape and duration and against changes in the width of cortical synchrony. The amplitude of PSF from a CM cell could be doubled by the presence of synchrony. It is proposed that, if a PSF has PWHM < 7 ms, this reliably indicates that the trigger is a CM cell projecting to the muscle whose EMG is averaged. In an analysis of experimental data where macaque motor cortical cells facilitated hand and forearm muscle EMG, 74% of PSFs fulfilled this criterion. The PWHM criterion could be applied to other STA studies in which it is important to exclude the effects of synchrony.

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.

Inspiratory drive and phase duration during carotid chemoreceptor stimulation in the cat: medullary neurone correlations.

1. This study addressed the hypothesis that there is a parallel processing of input from carotid chemoreceptors to brainstem neurones involved in inspiratory phase timing and control of inspiratory motor output amplitude. Data were from fifteen anaesthetized, bilaterally vagotomized, paralysed, artificially ventilated cats. Carotid chemoreceptors were stimulated by close arterial injection of 200 microliters of CO2-saturated saline solution. 2. Planar arrays of tungsten microelectrodes were used to monitor simultaneously up to twenty-two neurones in the nucleus tractus solitarii (NTS) and ventral respiratory group (VRG). Spike trains were analysed with two statistical tests of respiratory modulation, cycle-triggered histograms, peristimulus-time histograms, cumulative sum histograms and cross-correlograms. 3. In NTS, 16 of 26 neurones with respiratory and 12 of 27 without respiratory modulation changed firing rate during carotid chemoreceptor stimulation. In the VRG 72 of 112 respiratory and 14 of 48 non-respiratory neurones changed firing rate during stimulation. 4. The spike trains of 85 of 1276 pairs (6.7%) of cells exhibited short time scale correlations indicative of paucisynaptic interactions. Ten pairs of neurones were each composed of a rostral VRG phasic inspiratory neurone that responded to carotid chemoreceptor stimulation with a decline in firing rate and a caudal VRG phasic inspiratory neurone that increased its firing rate. Cross-correlograms from two of the pairs had features consistent with excitation of the caudal neurones by the rostral cells. A decrease in the duration of activity of the rostral VRG neurones was paralleled by the decrease in inspiratory time of phrenic nerve activity. Caudal VRG inspiratory neurones increased their activity as phrenic amplitude increased. Spike-triggered averages of all four neurones indicated post-spike facilitation of phrenic motoneurones. 5. The results support the hypothesis that unilateral stimulation of carotid chemoreceptors results in parallel actions. (a) Inhibition of rostral VRG I-Driver neurones decreases inspiratory duration. (b) Concurrent excitation of premotor VRG and dorsal respiratory group inspiratory neurones increases inspiratory drive to phrenic motoneurones. Other data suggest that responsive ipsilateral neurones act to regulate contralateral neurones.

Response patterns and postspike effects of premotor neurons in cervical spinal cord of behaving monkeys

Most of our information about spinal neurons has been derived from experiments with anesthetized or surgically. reduced preparations. To investigate these neurons under normal behavioral conditions, we recorded the activity of single afferent units in cervical dorsal root ganglia and of single interneurons in the cervical spinal cord of macaque monkeys, as they generated alternating flexion and extension torques about the wrist. Spike-triggered averages of rectified electromyographic activity were used to identify "premotor" (PreM) units associated with correlated postspike effects in active muscles. In addition to postspike effects, some spike-triggered averages showed early increases in average muscle activity, which were attributed to synchronous discharges in other PreM units. In recordings of peripheral afferents, 49% of the task-related dorsal root ganglia units produced postspike facilitation (PSF) of at least one forearm muscle, with a mean PSF latency of 5.8 +/- 0.3 ms (SE). The PSF amplitude was measured as the mean percent increase (MPI): the average increase of the PSF as a percentage of the prespike baseline mean. PreM afferent units produced PSF with an average MPI of 4.6 +/- 0.3%. In a study of cervical interneurons, about 13% (72/562) of the task-related cells showed postspike effects. These PreM interneurons had a mean PSF latency of 7.2 +/- 0.3 ms and a mean MPI of 4.6 +/- 0.2%. The MPI values for spinal neurons were similar to the MPIs reported for rubromotoneuronal and corticomotoneuronal cells. PreM neurons usually facilitated a subset of the coactivated muscles called the unit's "muscle field." The PreM afferents facilitated an average of 46% of the synergistically coactivated muscles, while PreM interneurons facilitated an average of 37%. These are comparable with the percentage of muscles facilitated by corticomotoneuronal (40%) and rubromotoneuronal (50%) cells. During the step-tracking task the monkeys generated ramp-and-hold torques about the wrist. The PreM afferents typically became active during either flexion or extension of the wrist, although a few were bidirectionally active. The most common response pattern in PreM afferents was a tonic discharge, followed by phasic and phasic-tonic discharge. The most common patterns exhibited by PreM interneurons were tonic and phasic-tonic responses. PreM afferent units began to discharge on average 51 +/- 13 ms before activation of their target muscle. This early onset supports our hypothesis that these PreM afferents arose from muscle spindles, which is also consistent with their short-latency PSF and their responses to perturbations that stretched their target muscles. The results reveal some salient differences between the discharge properties of dorsal root ganglia neurons, spinal interneurons, and supraspinal PreM cells in the motor cortex and red nucleus. All four PreM populations include tonic, phasic-tonic, and phasic cells, but in significantly different proportions. Most PreM afferents resembled corticomotoneuronal cells in being active only with their target muscles, unlike rubromotoneuronal cells and spinal PreM interneurons, which tended to exhibit more bidirectional discharges.

Functional linkages between motor cortical cells and elbow flexor muscles. Evidence for and characteristics of postspike facilitation.

1. Two monkeys (Macaca fascicularis) making high-level but submaximal isometric flexions of the elbow were investigated for the output effect of motor cortical cells on the electromyogram (EMG) activity of two main elbow flexors using the method of spike-triggered averaging of rectified EMGs (STAs). 2. Monkeys were trained to perform individual isometric contractions for > 2 s, and two series of > or = 20 contractions, the second series being at a greater force. EMG electrodes pairs were implanted in the biceps brachii and brachioradialis. A total of 257 cortical cells were found that discharged with the active and passive movements of the elbow. We examined the EMG postspike facilitations (PSFs) produced in either one or the two flexors for only those cells that discharged during the isometric contraction, and provoked PSFs in the two series of contractions. 3. The main characteristics of the EMG isometric contractions in the agonists were analyzed. Spectral analysis showed that the increases in the EMG median frequency with force stabilized at the force levels performed by monkeys. Cross correlation methods showed no cross talk between agonists. 4. The 26 selected cortical cells had a regular discharge frequency. Ten cells did not change frequency with a 22-30% force increase, 14 cells discharged at a higher frequency, and 2 cells discharged at a lower frequency. For single-cell frequencies of 5-65 Hz, interspike intervals < 10 ms were rare: the median and modal intervals were 20-30 ms. 5. The significance of PSFs with respect to the EMG background noise was estimated statistically. STAs from successive epochs under identical load conditions, and STAs performed at a distance from the trigger, showed that PSFs were authentic postspike effects and not sudden EMG changes synchronized by chance with the triggering cell. The features distinguishing PSF from secondary postspike EMG changes or coactivation and task-related effects were studied in simultaneous STAs of flexors and autocorrelogram of cortical spikes. 6. The magnitude of the PSF was expressed as the percent peak amplitude above the mean EMG baseline. The mean percent amplitude of the 90 PSFs produced in both muscles and series was 4.0 +/- 2.4% (mean +/- SD). There was no difference in the average amplitude of PSFs in the two flexors, although the baseline voltages in the biceps brachii were higher. Neither was there any significant change with force while the baseline level increased by 29 +/- 10%, indicating that the absolute PSF amplitude increased in the same proportion as baseline.(ABSTRACT TRUNCATED AT 400 WORDS)

Post‐spike facilitation of neck EMG by cat tectoreticulospinal neurones during orienting movements.

1. The activity of fourteen tectoreticulospinal neurones (TRSNs) was recorded intraaxonally in the caudal pons of alert cats during orienting movements towards visual stimuli. TRSN spikes were used to compute the spike-triggered average (STA) of rectified EMG of dorsal neck muscles. 2. Eight TRSNs for which 400-2532 spikes were available were analysed with the STA technique. When the STA was computed from all spikes, significant post-spike facilitation (PSF) was obtained for six of eighteen cell-muscle pairs investigated (5 TRSNs). The mean relative amplitude of PSFs was 7.4% (S.D. 3.7). The onset latencies ranged from 1.1 to 5.0 ms and mean duration was 11.4 +/- 3.1 ms (mean +/- S.D.). 3. Interspike interval distributions were unimodal, with modes between 2.7 and 12.7 ms. Spike trains of TRSNs that produced significant PSFs contained 5-13% of the interspike intervals < or = 5 ms and 22-37% of the intervals < or = 10 ms. To evaluate the contribution of short intervals to PSF, STAs were computed separately for spikes preceded by 'short' (< or = 5 or < or = 10 ms) and 'long' (> 5 or > 10 ms) intervals. 4. When computed from spikes preceded by 'long' intervals, PSF amplitudes were small (mean +/- S.D., 5.3 +/- 2.7%) and onset latencies measured by cusum ranged between 2.4 and 5.4 ms. This is longer than the estimated minimal latency of monosynaptic facilitatory effect on neck EMG (1.9-2.1 ms). 5. Relative amplitudes of PSF obtained with spikes preceded by 'short' intervals were much larger (mean +/- S.D., 14.8 +/- 7.4%), but cusums indicated negative latencies for four of six PSFs. The unrealistically short onset latencies could be accounted for by the summation of facilitation from the trigger spike with that of the preceding spikes. In four of five TRSNs a large increase of PSF amplitude (from 3.2 to 7.2 times the amplitude obtained from 'long' intervals) suggests the presence of frequency-dependent potentiation of synaptic transmission. 6. This study unequivocally demonstrates that some TRSNs produce significant post-spike facilitation of neck motoneurones. This facilitation could be mediated by monosynaptic tectomotoneuronal connections although a contribution by disynaptic connections cannot be definitively ruled out. The high instantaneous firing rates of TRSNs produce a potentiation of the otherwise weak facilitatory action of TRSNs that presumably contributes to a rapid recruitment of motoneurones during initiation of head orienting movements.

Characteristics of somatotopic organization and spontaneous neuronal activity in the region of the thalamic principal sensory nucleus in patients with spinal cord transection.

1. We explored the region of the principal sensory nucleus of thalamus (Vc) during stereotactic surgical procedures for treatment of patients with pain after spinal cord transection (n = 23). Receptive fields (RFs) of thalamic single neurons and locations of sensations evoked by stimulation (projected field, PF) were determined by standard methods. The cellular thalamic region where sensations were evoked at < 25 microA was termed the "region of Vc." The region of Vc in spinal patients was subdivided into different areas according to RF and PF locations. Areas that were distant from the representation of the anesthetic part of the body were termed "spinal control" areas, whereas those that were adjacent to or included in the representation of the area of absolute sensory loss were termed "border zone/anesthetic" areas. The region of Vc in movement disorder patients were termed the "control" area. 2. Border zone/anesthetic areas of thalamus often exhibited increased representations of the border of the anesthetic part of the body in comparison with the representation of the same parts of the body in control and spinal control areas. 3. In control and spinal control areas the locations of RFs and PFs were usually well matched. However, in border zone/anesthetic areas of the thalamus there was frequently a mismatch between the location of RFs and PFs (RF/PF mismatch). In border zone/anesthetic areas, RFs were often located on the border of the anesthetic part of the body whereas PFs were referred to anesthetic parts of the body. 4. Analysis of first- and higher-order properties of spontaneous neuronal activity revealed that spike trains could be classified into two groups with distinct patterns of activity. The R group (n = 49) was characterized by independence of sequential interspike intervals (ISIs), a Poisson distribution of ISIs, initially inhibitory or flat autocovariance function (acvf), and low level of high-frequency bursting. The O group (n = 26) was characterized by correlation of sequential ISIs, large sustained postspike facilitation on the acvf, and high prevalence of high-frequency bursting--all consistent with a bursting pattern of activity. A third group of spike trains (n = 17) had an initially inhibitory or flat acvf and a unimodal, positively shifted, ISI distribution that did not meet criteria for a Poisson distribution. 5. Spike trains in the R group were much more common in control and control spinal areas, whereas those in the O group were more common in border zone/anesthetic areas.(ABSTRACT TRUNCATED AT 400 WORDS)

Use of spike triggered averaging of muscle activity to quantify inputs to motoneuron pools

1. The goal of this study was to determine the extent to which postspike facilitation (PSpF) of electromyograms (EMGs) could be used to estimate the inputs to separate motoneuron pools, under conditions where there was wide variability in the parameters of muscle activity. These parameters included cancellation of motor unit action potentials (MUAPs), variations in EMG noise, and changes in MUAP amplitude and duration. A systematic series of computer simulations with increasing complexity were used to achieve this goal. The initial simulations (model I) included a premotoneuronal (PreM) cell connected to a single postsynaptic motoneuron (Mn), which in turn projected to a muscle. The next simulations (model II) included other target motoneurons with their efferents each projecting to separate muscles. The last simulations (model III) included more than one postsynaptic motoneuron per Mn-pool, as is the case in mammalian neuromuscular systems. 2. A sample simulation (model I) was performed to determine if the PreM-evoked effects were within physiologically observed values. A cross-correlogram (XC) calculated from a PreM cell and its target Mn, receiving a PreM-evoked excitatory postsynaptic potential (EPSP) of 0.5 mV, produced a XC peak area of 0.04 Mn-spikes/PreM-trigger. The PSpF of EMG activity evoked by this PreM cell had a mean percent increase of 4.6% (MPI = mean bin amplitude of PSpF above baseline/mean baseline level x 100). These XC and PSpF values were within the range of values previously obtained from animal experiments. 3. The magnitude of MUAP cancellation in the EMG was tested by calculating two spike-triggered averages (SpTAs) of EMGs from Mn-triggers (not PreM-triggers as in the other SpTAs): one using typical bipolar MUAPs and another using their rectified counterpart of only positive polarity to eliminate the possibility of MUAP cancellation. The PSpF calculated from bipolar spikes was 24.8% smaller than the one calculated using unipolar spikes. This cancellation could be greater or smaller depending on the state of parameters, such as the shape and number of MUAPs, that determine the probability of overlap between MUAP components of opposite polarity. All subsequent computer simulations used typical bipolar MUAPs. 4. A series of increasing motoneuron EPSP amplitudes were used to determine the relationship between PreM-Mn connection strength and PSpF area. A nearly perfect linear relationship between EPSP amplitude and PSpF area was obtained for SpTAs of rectified EMGs (r = 0.99). An equally linear relationship was obtained when averaging nonrectified EMGs (r = 0.99), but the smaller EPSPs or weaker synaptic connections were not detected.(ABSTRACT TRUNCATED AT 400 WORDS)