Modulation Of Discharge Rate Research Articles

Parkinsonism disrupts neuronal modulation in the pre-supplementary motor area during movement preparation.

Multiple studies suggest that Parkinson's disease (PD) is associated with changes in neuronal activity throughout the basal ganglia-thalamocortical motor circuit. There are limited electrophysiological data, however, describing how parkinsonism impacts neuronal activity in the pre-supplementary motor area (pre-SMA), an area in the medial frontal cortex involved in movement planning and motor control. In this study, single unit activity was recorded in the pre-SMA of two female non-human primates during a visually cued reaching task in both the naive and parkinsonian state using the 1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine (MPTP) model of parkinsonism. In the naive state neuronal discharge rates were dynamically modulated prior to the presentation of the instructional go-cue. In a subset of these modulated cells, the magnitude of modulation correlated linearly with reaction time (RT). In the parkinsonian state, however, modulation of discharge rates in the pre-SMA was disrupted and the predictive encoding of RT was significantly diminished. These findings add to our understanding of the role of pre-SMA in motor behavior and suggest that disrupted encoding in this cortical region contributes to the alteration of early preparatory and pre-movement processes present in Parkinson's disease.Significance statement Goal-directed movements, such as reaching for an object, necessitate temporal preparation and organization of information processing within the basal ganglia-thalamocortical motor network. Impaired movement in people with Parkinson's disease is thought to be the result of pathophysiological activity disrupting information flow within this network. This work provides neurophysiological evidence linking altered motor preplanning processes encoded in the neuronal firing pattern of pre-supplementary motor area (pre-SMA) cells to the pathogenesis of motor disturbances in parkinsonism.

Motor unit discharge rate modulation during isometric contractions to failure is intensity- and modality-dependent.

The physiological mechanisms determining the progressive decline in the maximal muscle torque production capacity during isometric contractions to task failure are known to depend on task demands. Task-specificity of the associated adjustments in motor unit discharge rate (MUDR), however, remains unclear. This study examined MUDR adjustments during different submaximal isometric knee extension tasks to failure. Participants performed a sustained and an intermittent task at 20% and 50% of maximal voluntary torque (MVT), respectively (Experiment 1). High-density surface EMG signals were recorded from vastus lateralis (VL) and medialis (VM) and decomposed into individual MU discharge timings, with the identified MUs tracked from recruitment to task failure. MUDR was quantified and normalised to intervals of 10% of contraction time (CT). MUDR of both muscles exhibited distinct modulation patterns in each task. During the 20% MVT sustained task, MUDR decreased until ∼50% CT, after which it gradually returned to baseline. Conversely, during the 50% MVT intermittent task, MUDR remained stable until ∼40-50% CT, after which it started to continually increase until task failure. To explore the effect of contraction intensity on the observed patterns, VL and VM MUDR was quantified during sustained contractions at 30% and 50% MVT (Experiment 2). During the 30% MVT sustained task, MUDR remained stable until ∼80-90% CT in both muscles, after which it continually increased until task failure. During the 50% MVT sustained task the increase in MUDR occurred earlier, after ∼70-80% CT. Our results suggest that adjustments in MUDR during submaximal isometric contractions to failure are contraction modality- and intensity-dependent. KEY POINTS: During prolonged muscle contractions a constant motor output can be maintained by recruitment of additional motor units and adjustments in their discharge rate. Whilst contraction-induced decrements in neuromuscular function are known to depend on task demands, task-specificity of motor unit discharge behaviour adjustments is still unclear. In this study, we tracked and compared discharge activity of several concurrently active motor units in the vastii muscles during different submaximal isometric knee extension tasks to failure, including intermittent vs. sustained contraction modalities performed in the same intensity domain (Experiment 1), and two sustained contractions performed at different intensities (Experiment 2). During each task, motor units modulated their discharge rate in a distinct, biphasic manner, with the modulation pattern depending on contraction intensity and modality. These results provide insight into motoneuronal adjustments during contraction tasks posing different demands on the neuromuscular system.

Motor unit modes in the calf muscles during a submaximal isometric contraction are changed by brief stretches.

The purpose of our study was to investigate the influence of a stretch intervention on the common modulation of discharge rate among motor units in the calf muscles during a submaximal isometric contraction. The current report comprises a computational analysis of a motor unit dataset that we published previously (Mazzo etal., 2021). Motor unit activity was recorded from the three main plantar flexor muscles while participants performed an isometric contraction at 10% of the maximal voluntary contraction force before and after each of two interventions. The interventions were a control task (standing balance) and static stretching of the plantar flexor muscles. A factorization analysis on the smoothed discharge rates of the motor units from all three muscles yielded three modes that were independent of the individual muscles. The composition of the modes was not changed by the standing-balance task, whereas the stretching exercise reduced the average correlation in the second mode and increased it in the third mode. A centroid analysis on the correlation values showed that most motor units were associated with two or three modes, which were presumed to indicate shared synaptic inputs. The percentage of motor units adjacent to the seven centroids changed after both interventions: Control intervention, mode 1 decreased and the shared mode 1+2 increased; stretch intervention, shared modes either decreased (1+2) or increased (1+3). These findings indicate that the neuromuscular adjustments during both interventions were sufficient to change the motor unit modes when the same task was performed after each intervention. KEY POINTS: Based on covariation of the discharge rates of motor units in the calf muscles during a submaximal isometric contraction, factor analysis was used to assign the correlated discharge trains to three motor unit modes. The motor unit modes were determined from the combined set of all identified motor units across the three muscles before and after each participant performed a control and a stretch intervention. The composition of the motor unit modes changed after the stretching exercise, but not after the control task (standing balance). A centroid analysis on the distribution of correlation values found that most motor units were associated with a shared centroid and this distribution, presumably reflecting shared synaptic input, changed after both interventions. Our results demonstrate how the distribution of multiple common synaptic inputs to the motor neurons innervating the plantar flexor muscles changes after a brief series of stretches.

1633-P: Type 1 Diabetes Induced Alterations in Motor Unit Discharge Rate

Type 1 diabetes (T1D) is a contributing factor in alterations in skeletal muscle health. It has been attributed to accelerated muscle ageing (Monaco et al., 2019), and the molecular alterations in the muscle are also associated with impaired adaptations to physical exercise (Minnock et al., 2022). However, there is no evidence about the consequences of this condition on the behaviour of individual motor units (MUs), despite them being the effectors of the contractile stimulus coming from the nervous system via modulation of their recruitment and discharge activity. This study aimed to investigate whether MUs discharge rate (DR) differs between people living with T1D compared with a control (CON) group of people living without diabetes. Eight people with T1D (4M/4F, 30.50 ± 3.66 years, HbA1c of 8.31 ± 2.40%) without comorbidities and 8 age and sex-matched CON participants undertook the study. MUs DR was evaluated from the vastus lateralis muscle during trapezoidal submaximal isometric contractions with High-Density Electromyography at 20, and 40% of the maximum voluntary contraction (MVC) and DR modulation was defined as the difference between DR at recruitment (i.e., first 3 MUs firings) and DR during the steady-state phase of the contraction. Groups were compared for maximum torque (182.06 ± 47.65 Nm T1D and 171.99 ± 101.97 Nm Non-D). MUs DR was lower for those with T1D at recruitment and during the steady-state phase, both during contractions at 20 and 40% MVC (p<0.05). MUs DR modulation was comparable between the two groups. In conclusion, reductions in MUs DR indicate initial alterations of MUs properties in people living with uncomplicated T1D when compared to CON counterparts. The reduced DR, however, was not accompanied by a reduced range of DR modulation (gain) and was not reflected in reduced force production. These findings suggest that initial alterations of MUs properties are present in people living with uncomplicated T1D and that these alterations are detectable before they become functionally manifest. Disclosure D.Minnock: Employee; Innovation Zed. G.Valli: None. R.Wu: None.

Respiration-related discharge of hyoglossus muscle motor units in the rat

Although respiratory muscle motor units have been studied during natural breathing, simultaneous measures of muscle force have never been obtained. Tongue retractor muscles, such as the hyoglossus (HG), play an important role in swallowing, licking, chewing, breathing, and, in humans, speech. The HG is phasically recruited during the inspiratory phase of the respiratory cycle. Moreover, in urethane anesthetized rats the drive to the HG waxes and wanes spontaneously, providing a unique opportunity to study motor unit firing patterns as the muscle is driven naturally by the central pattern generator for breathing. We recorded tongue retraction force, the whole HG muscle EMG and the activity of 38 HG motor units in spontaneously breathing anesthetized rats under low-force and high-force conditions. Activity in all cases was confined to the inspiratory phase of the respiratory cycle. Changes in the EMG were correlated significantly with corresponding changes in force, with the change in EMG able to predict 53-68% of the force variation. Mean and peak motor unit firing rates were greater under high-force conditions, although the magnitude of discharge rate modulation varied widely across the population. Changes in mean and peak firing rates were significantly correlated with the corresponding changes in force, but the correlations were weak (r(2) = 0.27 and 0.25, respectively). These data indicate that, during spontaneous breathing, recruitment of HG motor units plays a critical role in the control of muscle force, with firing rate modulation playing an important but lesser role.

Response variability of frontal eye field neurons modulates with sensory input and saccade preparation but not visual search salience

Discharge rate modulation of frontal eye field (FEF) neurons has been identified with a representation of visual search salience (physical conspicuity and behavioral relevance) and saccade preparation. We tested whether salience or saccade preparation are evident in the trial-to-trial variability of discharge rate. We quantified response variability via the Fano factor in FEF neurons recorded in monkeys performing efficient and inefficient visual search tasks. Response variability declined following stimulus presentation in most neurons, but despite clear discharge rate modulation, variability did not change with target salience. Instead, we found that response variability was modulated by stimulus luminance and the number of items in the visual field independently of attentional demands. Response variability declined to a minimum before saccade initiation, and presaccadic response variability was directionally tuned. In addition, response variability was correlated with the response time of memory-guided saccades. These results indicate that the trial-by-trial response variability of FEF neurons reflects saccade preparation and the strength of sensory input, but not visual search salience or attentional allocation.

Dependence of the paired motor unit analysis on motor unit discharge characteristics in the human tibialis anterior muscle

The paired motor unit analysis provides in vivo estimates of the magnitude of persistent inward currents (PIC) in human motoneurons by quantifying changes in the firing rate (Δ F) of an earlier recruited (reference) motor unit at the time of recruitment and derecruitment of a later recruited (test) motor unit. This study assessed the variability of Δ F estimates, and quantified the dependence of Δ F on the discharge characteristics of the motor units selected for analysis. Δ F was calculated for 158 pairs of motor units recorded from nine healthy individuals during repeated submaximal contractions of the tibialis anterior muscle. The mean (SD) Δ F was 3.7 (2.5) pps (range −4.2 to 8.9 pps). The median absolute difference in Δ F for the same motor unit pair across trials was 1.8 pps, and the minimal detectable change in Δ F required to exceed measurement error was 4.8 pps. Δ F was positively related to the amount of discharge rate modulation in the reference motor unit ( r 2 = 0.335; P < 0.001), and inversely related to the rate of increase in discharge rate ( r 2 = 0.125; P < 0.001). A quadratic function provided the best fit for relations between Δ F and the time between recruitment of the reference and test motor units ( r 2 = 0.229, P < 0.001), the duration of test motor unit activity ( r 2 = 0.110, P < 0.001), and the recruitment threshold of the test motor unit ( r 2 = 0.237, P < 0.001). Physiological and methodological contributions to the variability in Δ F estimates of PIC magnitude are discussed, and selection criteria to reduce these sources of variability are suggested for the paired motor unit analysis.

Discharge rate modulation of trapezius motor units differs for voluntary contractions and instructed muscle rest

This study examined discharge rate modulation at respiratory (0-0.5 Hz) and beta (16-32 Hz) frequencies in trapezius motor units active during voluntary contractions and during periods of instructed rest under conditions of low and high psychosocial stress. In separate sessions, single motor unit activity was recorded from the trapezius muscle of healthy women during low-intensity voluntary contractions and during periods of instructed muscle rest that followed voluntary contractions. The level of psychosocial stress during periods of instructed muscle rest was manipulated using a verbal math task combined with social evaluative threat which increased perceived anxiety, heart rate, and blood pressure (P ≤ 0.002). Discharge rate modulation was quantified by the mean power of motor unit discharge rate profiles within frequency bands of interest. Under low stress conditions, motor units active during instructed rest had greater power at 0-0.5 Hz (P = 0.002) and less power at 16-32 Hz (P = 0.009) compared to those active during voluntary contraction. Exposure to the stressor increased the amount of motor unit activity during instructed rest (P = 0.021) but did not alter the power of discharge rate modulation at 0-0.5 Hz (P = 0.391) or 16-32 Hz (P = 0.089). These results indicate that sustained motor unit activity during periods of instructed muscle rest has a lesser contribution from inputs at beta frequencies and a greater contribution from inputs at respiratory frequencies than present during low-intensity voluntary contractions. Furthermore, increases in motor unit activity when exposed to stressors during periods of instructed rest are not caused by changes in inputs at respiratory or beta frequencies.

Cooperation and Competition among Frontal Eye Field Neurons during Visual Target Selection

The role of spike rate versus timing codes in visual target selection is unclear. We simultaneously recorded activity from multiple frontal eye field neurons and asked whether they interacted to select targets from distractors during visual search. When both neurons in a pair selected the target and had overlapping receptive fields (RFs), they cooperated more than when one or neither neuron in the pair selected the target, measured by positive spike timing correlations using joint peristimulus time histogram analysis. The amount of cooperation depended on the location of the search target: it was higher when the target was inside both neurons' RFs than when it was inside one RF but not the other, or outside both RFs. Elevated spike timing coincidences occurred at the time of attentional selection of the target as measured by average modulation of discharge rates. We observed competition among neurons with spatially non-overlapping RFs, measured by negative spike timing correlations. Thus, we provide evidence for dynamic and task-dependent cooperation and competition among frontal eye field neurons during visual target selection.

Discharge Behaviors of Trapezius Motor Units During Exposure to Low and High Levels of Acute Psychosocial Stress

This study investigated the effects of acute psychosocial stress on trapezius single motor unit discharge behaviors. Twenty-one healthy women performed feedback-controlled isometric contractions under conditions of low and high psychosocial stress in the same experimental session. Psychosocial stress was manipulated using a verbal math task combined with social evaluative threat that significantly increased perceived anxiety, heart rate, and blood pressure (P < 0.001). Motor unit discharge behaviors including the threshold and discharge rate at recruitment [7.7% (5.7%) maximal voluntary isometric contraction and 7.3 pulses per second (pps) (6.8 pps), P > 0.121, N = 103] and derecruitment [6.0% (4.4%) maximal voluntary isometric contraction and 6.5 pps (4.1 pps), P > 0.223, N = 99], the mean [11.3 pps (2.3 pps), P = 0.309, N = 106] and variability [2.5 pps (0.91 pps), P = 0.958, N = 106] of discharge rate, and the proportion of motor units exhibiting double discharges (21%, P = 0.446) did not change across stress conditions. Discharge rate modulation with changes in contraction intensity was highly variable and similar across stress conditions (P > 0.308, N = 89). Rate-rate modulation of concurrently active motor units was also highly variable (r = -0.84 to 1.00, N = 75). Estimates of DeltaF for motor unit pairs with rate-rate modulation >or=0.7 were positive and similar across stress conditions [4.7 pps (2.0 pps), P = 0.405, N = 16]. The results indicate that acute psychosocial stress does not alter trapezius motor unit discharge behaviors during a precisely controlled motor task in healthy women.

Effect of pain on the modulation in discharge rate of sternocleidomastoid motor units with force direction

To compare the behavior of sternocleidomastoid motor units of patients with chronic neck pain and healthy controls. Nine women (age, 40.4+/-3.5 yr) with chronic neck pain and nine age- and gender-matched healthy controls participated. Surface and intramuscular EMG were recorded from the sternocleidomastoid muscle bilaterally as subjects performed isometric contractions of 10-s duration in the horizontal plane at a force of 15 N in eight directions (0-360 degrees ; 45 degrees intervals) and isometric contractions at 15 and 30 N force with continuous change in force direction in the range 0-360 degrees . Motor unit behavior was monitored during the 10-s contractions and the subsequent resting periods. The mean motor unit discharge rate depended on the direction of force in the control subjects (P<0.05) but not in the patients. Moreover, in three of the nine patients, but in none of the controls, single motor unit activity continued for 8.1+/-6.1s upon completion of the contraction. The surface EMG amplitude during the circular contraction at 15N was greater for the patients (43.5+/-54.2 microV) compared to controls (16.9+/-14.9 microV; P<0.05). The modulation in discharge rate of individual motor units with force direction is reduced in the sternocleidomastoid muscle in patients with neck pain, with some patients showing prolonged motor unit activity when they were instructed to rest. These observations suggest that chronic neck pain affects the change in neural drive to muscles with force direction.

Role of supplementary eye field in saccade initiation: executive, not direct, control.

The goal of this study was to determine whether the activity of neurons in the supplementary eye field (SEF) is sufficient to control saccade initiation in macaque monkeys performing a saccade countermanding (stop signal) task. As previously observed, many neurons in the SEF increase the discharge rate before saccade initiation. However, when saccades are canceled in response to a stop signal, effectively no neurons with presaccadic activity display discharge rate modulation early enough to contribute to saccade cancellation. Moreover, SEF neurons do not exhibit a specific threshold discharge rate that could trigger saccade initiation. Yet, we observed more subtle relations between SEF activation and saccade production. The activity of numerous SEF neurons was correlated with response time and varied with sequential adjustments in response latency. Trials in which monkeys canceled or produced a saccade in a stop signal trial were distinguished by a modest difference in discharge rate of these SEF neurons before stop signal or target presentation. These findings indicate that neurons in the SEF, in contrast to counterparts in the frontal eye field and superior colliculus, do not contribute directly and immediately to the initiation of visually guided saccades. However the SEF may proactively regulate saccade production by biasing the balance between gaze-holding and gaze-shifting based on prior performance and anticipated task requirements.

Comments on Point:Counterpoint: Spectral properties of the surface EMG can characterize/do not provide information about motor unit recruitment strategies and muscle fiber type

to the editor: Motor unit activity can be characterized either with direct physiological measurements or with indirect estimates that are based on the association of another parameter with the physiology. Key physiological properties include contractile characteristics, discharge rate modulation,

Effect of muscle‐fiber velocity recovery function on motor unit action potential properties in voluntary contractions

Measurements of muscle-fiber conduction velocity during voluntary contractions have been used in the diagnosis of neuromuscular diseases. However, the velocity of propagation of action potentials depends on the interspike interval of activation due to the velocity recovery function (VRF) of muscle fibers. The comparison of muscle-fiber conduction velocity estimates between individuals may thus be influenced by differences in motor unit discharge rate. This study investigates action potential properties of motor units of the sternocleidomastoid muscle during voluntary modulation of discharge rate with the purpose of assessing the effect of the VRF on motor unit properties in voluntary contractions. Nineteen healthy men trained to control a target motor unit with feedback of surface multichannel electromyographic (EMG) signals. The subjects performed three 30-s contractions of cervical flexion/rotation modulating the discharge rate of the target motor unit from 6.6 +/- 1.6 pps to 28.0 +/- 6.4 pps. Action potential conduction velocity was correlated to instantaneous discharge rate (R = 0.38 +/- 0.21). Action potential conduction velocity, peak-to-peak amplitude, and duration varied between minimum and maximum discharge rate (P < 0.01; percent change 12.3 +/- 5.0, -11.8 +/- 9.9, and -12.9 +/- 7.3). Thus, the properties of surface motor unit action potentials vary with modulation of discharge rate. This has implications for the use of conduction velocity values measured during voluntary contractions to differentiate patient populations from healthy individuals and for the development of normative data.

Motor units in cranial and caudal regions of the upper trapezius muscle have different discharge rates during brief static contractions

To compare the discharge patterns of motor unit populations from different locations within the upper trapezius muscle during brief submaximal constant-force contractions. Intramuscular and surface electromyographic (EMG) signals were collected from three sites of the right upper trapezius muscle distributed along the cranial-caudal direction in 11 volunteers during 10 s shoulder abduction at 25% of the maximum voluntary force. A total of 38 motor units were identified at the cranial location, 36 from the middle location and 17 from the caudal location. Initial discharge rate was greatest at the caudal location (P < 0.05; mean +/- SD, cranial: 16.7 +/- 3.6 pps, middle: 16.9 +/- 4.0 pps, caudal: 19.2 +/- 3.3 pps). Discharge rate decreased during the contraction for the most caudal location only (P < 0.05). Initial estimates of surface EMG root mean square values were highest at the most caudal location (P < 0.05; cranial: 32.3 +/- 20.9 microV, middle: 41.3 +/- 21.0 microV, caudal: 51.6 +/- 23.6 microV). This study demonstrates non-uniformity of motor unit discharge within the upper trapezius muscle during a brief submaximal constant-force contraction. Location-dependent modulation of discharge rate may reflect spatial dependency in the control of motor units necessary for the development and maintenance of force output.

Specific modulation of motor unit discharge for a similar change in fascicle length during shortening and lengthening contractions in humans

This study examines the effect of a change in fascicle length on motor unit recruitment and discharge rate in the human tibialis anterior during shortening and lengthening contractions that involved a similar change in torque. The dorsiflexor torque and the surface and intramuscular electromyograms (EMGs) from the tibialis anterior were recorded in eight subjects. The behaviour of the same motor unit (n=63) was compared during submaximal shortening and lengthening contractions performed at a constant velocity (10 deg s-1) with the dorsiflexor muscles over a 20 deg range of motion around the ankle neutral position. Muscle fascicle length was measured non-invasively using ultrasonography. Motor units that were active during a shortening contraction were always active during the subsequent lengthening contraction. Furthermore, additional motor units (n=18) of higher force threshold that were recruited during the shortening contraction to maintain the required torque were derecruited first during the following lengthening contraction. Although the change in fascicle length was linear (r2>0.99), and similar for both shortening and lengthening contractions, modulation of discharge rate differed during the two contractions. Compared with an initial isometric contraction at short (11.9+/-2.4 Hz) or long (11.7+/-2.2 Hz) muscle length, discharge rate increased only slightly and stayed nearly constant throughout the lengthening contraction (12.6+/-2.0 Hz; P<0.05) whereas it augmented progressively and more substantially during the shortening contraction, reaching 14.5+/-2.5 Hz (P<0.001) at the end of the movement. In conclusion, these observations indicate a clear difference in motor unit discharge rate modulation with no change in their recruitment order between shortening and lengthening contractions when performed with a similar change in muscle fascicle length and torque.

Reinnervation of motor units in intrinsic muscles of a transplanted hand

Functional recovery of transplanted hand can be evaluated clinically but until now there has been no direct assessment of muscle control. In October 2000 we transplanted the right hand of a brain-dead man aged 43 onto a man aged 35 who had lost his right dominant hand 22 years before. Starting from day 205 after the transplant, multi-channel surface electromyographic (EMG) signals were recorded from intrinsic muscles of the transplanted hand in order to assess their degree of reinnervation. Eleven months post-operatively, the first motor unit action potential train was detected from the abductor digiti minimi. One month later, also the abductor pollicis brevis and the opponens pollicis muscles showed motor unit activity, while, after 15 and 24 months, the first dorsal interosseous and the first lumbricalis muscles, respectively, showed activation of their first motor units. An increase in the number of active motor units was observed after the first signs of reinnervation, although the process was rather slow. In sustained maximal contractions, the motor unit discharge rate decreased from (mean ± S.D.) 34.0 ± 6.7 pps to 23.4 ± 5.1 pps in 60 s for the abductor digiti minimi, although the subject was verbally encouraged to maintain a maximal activation. Moreover, the subject was able to perform basic control tasks involving voluntary modulation of motor unit discharge rate. With a visual feedback, he could increase discharge rate of the abductor digiti minimi approximately linearly over time, from 13.4 ± 6.7 pps to 32.5 ± 11.2 pps in 60 s. In conclusion, we showed reinnervation of single motor units in a transplanted hand after 22 years of denervation. Moreover, voluntary modulation of discharge rates of these motor units could be performed since the first sign of reinnervation.

Enhanced motor unit rate coding with improvements in a force-matching task

These data describe improved modulation of discharge rates (rate coding) of first dorsal interosseous motor units throughout the acquisition of a complex force-matching skill involving isometric index finger abduction. In each of 15 consecutive trials, subjects attempted to match their force to a trajectory consisting of the sum of two sine waves (0.15 and 0.5 Hz) and random oscillations (overall mean force level ~20% MVC). Reductions in root-mean-square (RMS) error of each subject’s force relative to the trajectory indicated substantial improvements in force-matching ability ( F=33.8, p<0.001). With the acquisition of this new skill, there was increased amplitude modulation of muscular force near both dominant frequencies of the force-matching trajectory ( F=10.6, p=0.008). The standard deviation and coefficient of variation of motor unit inter-spike intervals both decreased with improved performance indicating a general reduction in the amplitude of firing rate modulations (SD: F=18.69, p=0.001; CV: F=43.6, p<0.001). After skill acquisition, there was decreased firing rate modulation outside of the two dominant frequencies and increased amplitude of firing rate modulation at the higher of the two dominant frequencies (0.5 Hz, F=8.23, p=0.015). These findings indicate that improved precision of rate coding was a contributor to the acquisition of the new force-matching task. That the change in rate coding was frequency dependent suggests that factors other than frequency coding may contribute to the improved force matching at 0.15 Hz.

Context dependency in the globus pallidus internal segment during targeted arm movements.

Extracellular discharges from single neurons in the internal segment of the globus pallidus (GPi) were recorded and analyzed for rate changes associated with visually guided forearm rotations to four different targets. We sought to examine how GPi neurons contribute to movement preparation and execution. Unit discharge from 108 GPi neurons recorded in 35 electrode penetrations was aligned to the time of various behavioral events, including the onset of cued and return movements. In total, 39 of 108 GPi neurons (36%) were task-modulated, demonstrating statistically significant changes in discharge rate at various times between the presentation of visual cues and movement generation. Most often, strong modulation in discharge rate occurred selectively during either the cued (n = 32) or return (n = 2) phases of the task, although a few neurons (n = 5) were well-modulated during both movement phases. Of the 34 neurons that were modulated exclusively during cued or return movements, 50% (n = 17) were modulated similarly in association with movements to any target. The remaining 17 neurons exhibited considerable diversity in their discharge properties associated with movements to each target. Cued phases of behavior were always rewarded if executed correctly, whereas return phases were never rewarded. Overall, these data reveal that many GPi neurons discharged in a context-dependent manner, being modulated during cued, rewarded movements, but not during similar self-paced, unrewarded movements. When considered in the light of other observations, the context-dependence we have observed seems likely to be influenced by the animal's expectation of reward.

Spontaneous discharge and response characteristics of central otolith neurons of rats during postnatal development.

To study the developmental profile of otolith-related vestibular nuclear neurons, their spontaneous activities and response dynamics were examined in decerebrate rats aged seven, 14, 21 and 84 (adult) days. Extracellular recordings were performed in the lateral and descending vestibular nucleus of animals held at the stationary position in the earth-horizontal or subjected to constant velocity off-vertical axis rotation, which selectively stimulates the otolith receptors. All neurons displayed sinusoidal position-dependent modulation in discharge rate, indicating their capability in coding spatial information during low-frequency head movement. Some neurons showed a full-cycle response to off-vertical axis rotation (non-clipped), while other neurons were silenced in discharge during parts of each rotary cycle (clipped). In seven-day-old rats, three-quarters of the responsive neurons sampled were clipped and the proportion progressively decreased to less than one-quarter in adult rats. In each age group, the clipped neurons discharged in approximately 60% of the stimulus cycle. Response gains of the neurons increased with age, reaching a plateau from 21 days of age for clipped neurons and 14 days for non-clipped neurons. The clipped neurons demonstrated higher response gains than the non-clipped neurons at or beyond 21 days of age. Spontaneous activities of the neurons at the stationary and earth-horizontal positions were analysed in relation to their response gains; a positive correlation was observed from 14 days of age onwards. Both types of neurons showed progressive increase in spontaneous activity as the rats matured, though the clipped neurons exhibited significantly lower resting rates than the non-clipped neurons at each of the age groups studied. Some neurons that responded to off-vertical axis rotation were not spontaneously active at the stationary position, but the proportion of these decreased significantly with age. The coefficient of variation of each age group showed a bimodal distribution, thereby allowing spontaneously active neurons to be assigned as regular or irregular. Though the vast majority of both the clipped and non-clipped neurons showed irregular discharge patterns at seven days of age, the overall population became more regular as the rats matured. Irregular neurons of young rats exhibited phase-stable and phase-shift responses, while those of older rats showed only the phase-stable response. This distinction was not observed amongst regular neurons over the ages studied. Our results reveal features of central otolith neurons that can be taken as signs of maturation during the first three postnatal weeks. These neuronal features provide the framework for the analysis of behaviours mediated by the otolith system during postnatal maturation.