Recruitment Order Of Motor Units Research Articles

Functional consequences of motor unit recruitment order reversals following spinal cord transection in cat

Motoneuron recruitment order determinations were made for acute, 2-week chronic, and 3-month chronic spinal cats by comparing cutaneous nerve stimulation thresholds for evoking single unit tibialis anterior (TA) electromyogram (EMG) spikes of different sizes. Recruitment order was largely (∼80%) orderly (small spikes recruited at lower stimulus intensities than large spikes) in acute and 3-month chronic spinal animals. However, in 2-week chronic spinal animals recruitment order was reversed, with large units more often recruited at lower stimulus intensities than small units (∼65%). Morphological analyses of TA muscle fibers suggested that fiber size changes were unlikely to account for the dramatic alterations in recruitment order results of the 2-week chronic spinal animals. Additional studies suggested that the recruitment order reversal in the 2-week chronic animals coincided with an enhanced reflex neural output (increased recruitment or reflex gain) for the flexion reflex which compensated for disuse atrophy related decreases in flexor muscle force generation capability in these animals. The data from 2-week chronic spinal animals represent a functionally significant example of deviation from the normal size principle of motoneuron recruitment order as the corresponding reflex gain increases can enhance the rapidity of motor function recovery (standing, locomotion) following spinal injury.

Motor unit behavior during clonus

Medial gastrocnemius surface electromyographic activity and intramuscular electromyographic activity were recorded from six individuals with chronic cervical spinal cord injury to document the recruitment order of motor units during clonus. Four subjects induced clonus that lasted up to 30 s while two subjects induced clonus that they actively stopped after 1 min. Mean clonus frequency in different subjects ranged from 4.7 to 7.0 Hz. Most of the 166 motor units recorded during clonus (98%) fired once during each contraction but at slightly different times during each cycle. Other motor units fired during some clonus cycles (1%) or in bursts (1%). When 59 pairs of units were monitored over consecutive clonus cycles (n = 5-89 cycles), only 8 pairs of units altered their recruitment order in some cycles. Recruitment reversals only occurred in units that fired close together in the clonus cycle. These data demonstrate that orderly motor unit recruitment occurs during involuntary contractions of muscles paralyzed chronically by cervical spinal cord injury, providing further support for the importance of spinal mechanisms in the control of human motor unit behavior.

Motor unit recruitment during vertebrate locomotion

AbstractVertebrate skeletal muscles act across joints and produce segmental accelerations and therefore animal movement when they contract. Different muscles and different motor units vary in their mechanical contractile properties. Early studies on motor unit recruitment demonstrated orderly recruitment of motor units from the slowest to the fastest during a graded contraction. However, many subsequent studies illustrate conditions when alternative recruitment strategies may exist. Motor unit recruitment during locomotion is thus multifactorial and more complex than typically thought.Different types of motor unit vary in their mechanical properties, including rates of force activation and deactivation, maximum unloaded shortening velocities and the shortening velocities at which maximum mechanical power output and maximum mechanical efficiency occur. In short, it would make mechanical sense to perform fast activities with the faster motor units and slow activities with the slower motor units. However, determining patterns of motor unit recruitment during locomotion has presented experimental challenges.Comparisons between distinct muscles have shown that fast fish swimming and the cat paw shake are activities which employ predominantly the fast and not the slower muscle. Glycogen depletion studies have showed that jumping in the bushbaby uses fast without slow motor units within the vastus lateralis and gastrocnemius muscles. Studies in man show that differential recruitment of the different types of muscle fibre occurs at different times within each running stride. It is suggested that vertebrates may have a strategy of recruiting the motor units that are most mechanically suited for the different locomotor demands. However, we have much to learn about motor recruitment patterns during locomotion.

Recruitment order of motor units in human vastus lateralis muscle is maintained during fatiguing contractions.

Motor-unit firing patterns were studied in the vastus lateralis muscle of five healthy young men [21.4 +/- 0.9 (SD) yr] during a series of isometric knee extensions performed to exhaustion. Each contraction was held at a constant torque level, set to 20% of the maximal voluntary contraction at the beginning of the experiment. Electromyographic signals, recorded via a quadrifilar fine wire electrode, were processed with the precision decomposition technique to identify the firing times of individual motor units. In repeat experiments, whole-muscle mechanical properties were measured during the fatigue protocol using electrical stimulation. The main findings were a monotonic decrease in the recruitment threshold of all motor units and the progressive recruitment of new units, all without a change of the recruitment order. Motor units from the same subject showed a similar time course of threshold decline, but this decline varied among subjects (mean threshold decrease ranged from 23 to 73%). The mean threshold decline was linearly correlated (R2 >or= 0.96) with a decline in the elicited peak tetanic torque. In summary, the maintenance of recruitment order during fatigue strongly supports the notion that the observed common recruitment adaptations were a direct consequence of an increased excitatory drive to the motor unit pool. It is suggested that the increased central drive was necessary to compensate for the loss in force output from motor units whose muscle fibers were actively contracting. We therefore conclude that the control scheme of motor-unit recruitment remains invariant during fatigue at least in relatively large muscles performing submaximal isometric contractions.

Use of surface EMG for studying motor unit recruitment during isometric linear force ramp

The aim of this work was to demonstrate the rank order of motor unit (MU) recruitment by surface EMG based on a Laplacian detection technique and to document the MU features at their recruitment threshold. Surface EMG signals were recorded on the biceps brachii of 10 healthy subjects during linear force ramps. When achievable, the signals were decomposed into MU action potential (MUAP) trains. MU inter-pulse interval (IPI), conduction velocity (MUCV) and amplitude were estimated on the first 12 MUAPs of each detectable train in order to characterize the MU features at their firing onset. A strong correlation was found between MU recruitment threshold and IPI, MUCV, and amplitude, showing that the size principle can be demonstrated by a fully non-invasive EMG technique. However, signal decomposition was not possible on seven subjects due to the effects of the volume conductor when the skinfold thickness was too large. When requirements for an optimal detection of MUAP trains are met, surface EMG may be used to improve our understanding of MU activity.

Motor unit identification in two neighboring recording positions of the human trapezius muscle during prolonged computer work.

Work-related shoulder-neck pain is a major health risk in computer operators. To understand the physiological mechanisms behind the development of these disorders, EMG recordings of some minutes up to several hours must be accurately decomposed. For this reason we developed EMG-LODEC, an automatic decomposition software program, especially designed for multi-channel long-term recordings of signals detected during slight muscle movements. The subjects executed a 30-min computer task to simulate real work conditions while working at an ergonomically designed workstation. Six-channel intramuscular EMG signals were recorded from two positions of the upper trapezius muscle. The EMG signals were decomposed into individual motor unit action potential trains using EMG-LODEC. The study design enabled us first to study the dependence of intramuscular analysis on the insertion points and second to test the accuracy of the decomposition technique under laboratory conditions during a real experiment. The two positions yielded 887 motor units--452 located in position 1 and 435 in position 2. Although the numbers of detected action potentials were strongly correlated between the two insertion positions, different motor units were mostly recorded. In particular, the detection of continuously active motor units is specific for the selected insertion points and may not be representative of a muscle, not even for parts with common functions. The approach for the quantitative evaluation of the decomposition technique was to independently decompose two signals that were simultaneously detected by separate sets of wire electrodes placed close to each other in the muscle. Common trains discovered in each signal were compared for consistency. A cross-correlation analysis was performed to find corresponding motor unit pairs that were concurrently active. Concurrently active motor units were found in six subjects. For these motor units the extent of simultaneous occurrence of motor unit action potentials between the two positions ranged from 23% to 78% depending on the distinction of the single motor units and the number of superimposed motor unit action potentials. High concordance was seen in 3 out of the 15 motor unit pairs. Based on the results, EMG-LODEC is capable of providing reliable decompositions with satisfying accuracy and reasonable processing time. EMG-LODEC is suitable for the study of motor unit discharge patterns and recruitment order in subjects with and without musculoskeletal pain during long-term measurements to study work-related musculoskeletal disorders.

A nerve stimulation method to selectively recruit smaller motor-units in rat skeletal muscle

Electrical stimulation of peripheral nerve results in a motor-unit recruitment order opposite to that attained by natural neural control, i.e. from large, fast-fatiguing to progressively smaller, fatigue-resistant motor-units. Yet animal studies involving physiological exercise protocols of low intensity and long duration require minimal fatigue. The present study sought to apply a nerve stimulation method to selectively recruit smaller motor-units in rat skeletal muscle. Two pulse generators were used, independently supplying short supramaximal cathodal stimulating pulses (0.5 ms) and long subthreshold cathodal inactivating pulses (1.5 s) to the sciatic nerve. Propagation of action potentials was selectively blocked in nerve fibres of different diameter by adjusting the strength of the inactivating current. A tensile-testing machine was used to gauge isometric muscle force of the plantaris and both heads of the gastrocnemius muscle. The order of motor-unit recruitment was estimated from twitch characteristics, i.e. peak force and relaxation time. The results showed prolonged relaxation at lower twitch peak forces as the intensity of the inactivating current increased, indicating a reduction of the number of large motor-units to force production. It is shown that the nerve stimulation method described is effective in mimicking physiological muscle control.

Morphology and physiology of masticatory muscle motor units.

Motor unit territories in masticatory muscles appear to be smaller than territories in limb muscles, and this would suggest a more localized organization of motor control in masticatory muscles. Motor unit cross-sectional areas show a wide range of values, which explains the large variability of motor unit force output. The proportion of motor unit muscle fibers containing more than one myosin heavy-chain (MHC) isoform is considerably larger in masticatory muscles than in limb and trunk muscles. This explains the continuous range of contraction speeds found in masticatory muscle motor units. Hence, in masticatory muscles, a finer gradation of force and contraction speeds is possible than in limb and in trunk muscles. The proportion of slow-type motor units is relatively large in deep and anterior masticatory muscle regions, whereas more fast-type units are more common in the superficial and posterior muscle regions. Muscle portions with a high proportion of slow-type motor units are better equipped for a finer control of muscle force and a larger resistance to fatigue during chewing and biting than muscle portions with a high proportion of fast units. For the force modulation, masticatory muscles rely mostly on recruitment gradation at low force levels and on rate gradation at high force levels. Henneman's principle of an orderly recruitment of motor units has also been reported for various masticatory muscles. The presence of localized motor unit territories and task-specific motor unit activity facilitates differential control of separate muscle portions. This gives the masticatory muscles the capacity of producing a large diversity of mechanical actions. In this review, the properties of masticatory muscle motor units are discussed.

Muscular rest and gap frequency as EMG measures of physical exposure: the impact of work tasks and individual related

Owing to an orderly recruitment of motor units, low threshold type I fibres are presumed to be vulnerable in contractions of long duration. To study load on these fibres muscular rest was registered as the time fraction of electromyographic (EMG) activity below a threshold. Moreover, the frequency of periods with muscular rest, EMG gaps, was derived, since a low gap frequency has been shown to be a risk factor for musculoskeletal disorders. Trapezius EMG was registered in 24 female hospital cleaners, 21 female office workers and 13 male office workers during one working day. Cleaners have a high risk of neck/shoulder pain and had much less muscular rest than office workers measured as a percentage of total registered time (median value= 1.5%, range= 0.2-13% vs. median value= 12%, range= 0.0-32%, respectively). Gap frequency showed no difference between the two occupational groups. Both measures displayed a wide inter-individual variation. For the cleaners, some of the variance was explained by body mass index (BMI) and age, with lower values of muscular rest for older subjects with a high BMI. Among the office workers, low values of muscular rest and a high gap frequency were registered in subjects with a low subjective muscular tension tendency. Gender, strength, smoking, job strain, employment time and musculoskeletal symptoms had no impact on either EMG measure.

Human muscle activity related to non-biomechanical factors in the workplace.

This paper presents current knowledge on low-level, long-lasting work-related muscle activity, focusing on the shoulder and the upper part of the trapezius muscle, and on mental, rather than biomechanical reasons for the muscle activation. The paper identifies three sources of vocational muscle activity: the biomechanical need for force production in order to perform movements or maintain postures against the force of gravity, the biomechanical need to stabilise body parts as a reference for performing movements and securing a stable visual field, and finally, muscle activity without obvious biomechanical purposes. This last category has been labelled non-biomechanical muscle activity in this review. Non-biomechanical muscle activity is related to the mental load, the emotional load and the individual characteristics of the subject, and is identified as having a low-level and a low second-to-second variability, resembling a static muscle contraction. Recent research has indicated that the size principle for motor unit recruitment order puts a strain on a limited number of low-threshold motor units which might be heavily taxed despite the overall low level of this muscle activity. However, the paper also cites a recent report showing that motor unit substitution may occur in prolonged low-level muscle activation (longer than a few minutes). Evaluations of muscle load at work usually omit the possibility of extra muscle activation due to nonbiomechanical factors, and thus may often give estimates of the muscle load that are too low, or misinterpret nonbiomechanical muscle activity as biomechanical muscle load.

Muscle recruitment through electrical stimulation of the lumbo-sacral spinal cord.

The goal of this study was to determine the feasibility of producing graded muscle contraction in individual muscles or muscle groups by electrically stimulating motor neurons in the lumbo-sacral spinal cord. Recruitment curves were obtained for quadriceps, tibialis anterior and triceps surae/plantaris by stimulating their activation pools in the ventral horn of the feline spinal cord. Mean twitch times-to-peak for quadriceps, tibialis anterior and triceps surae/plantaris were 33.0, 41.0, and 36.0 ms, respectively. Twitch duration as a function of stimulus strength demonstrated a mixed motor unit recruitment order, distinctively different from the inverse recruitment order exhibited by conventional methods of electrical stimulation of peripheral nerve. The recruitment curve slopes (expressed as a percentage of maximum force per nanocurrent of delivered charge) were shallow: 7.9 for quadriceps, 2.6 for tibialis anterior and 8.5 for triceps surae/plantaris. These results show that graded control of force in individual muscles or muscle groups can be obtained through spinal cord stimulation, and suggest that spinal cord stimulation could be used for functional neuromuscular stimulation applications.

Electromyographic evidence for functional heterogeneity in the inferior head of the human lateral pterygoid muscle: a preliminary multi-unit study

Objective: Functional heterogeneity, i.e. regional or selective activation of subpopulations of fibres within a muscle, has been described in some jaw and limb muscles. Each head of the lateral pterygoid muscle may also be functionally heterogeneous. The aims of this investigation were to develop a technique to test this hypothesis, and to use this technique to determine whether there is any multi-unit electromyographic (EMG) evidence for functional heterogeneity within the inferior head of the lateral pterygoid (IHLP). Methods: In 3 human subjects without craniomandibular disorders, recordings were made of condylar movement and multi-unit EMG activity from two sites in the IHLP during repeated trials of a contralateral ( n=21) and a protrusive ( n=26) jaw movement. The recording sites within IHLP were approached extraorally (labelled IHLP-extra) and were verified by computer tomography (CT); the other (IHLP-intra) were from sites in IHLP approached intraorally. Results: In each subject, the time of occurrence of the peak filtered signal from IHLP-extra was significantly different ( P<0.05) from IHLP-intra for all protrusion trials but not for contralateral trials. Conclusions: The data suggest a task-dependent change in motor unit recruitment order within IHLP and that IHLP is functionally heterogeneous.

Force feedback control of motor unit recruitment in isometric muscle

The use of simple force feedback in an isometric muscle control system utilizing orderly recruitment of motor units is explored. Cat medial gastrocnemius motor units were stimulated with and without simple force feedback gain ranging from 0.7 to 0.9. Ramp, triangular, staircase, sinusoidal and bandwidth-limited pseudo-random input recruitment signals were used to study tracking accuracy through linear correlation in ramp and triangular signals, cross correlation in sinusoidal and random signals, and rise time and steady state error in staircase signals. Dramatic improvements were found in most tested tracking variables with the use of feedback; squared correlation coefficients increased from a mean of 0.93 to 0.99 for ramp signals and from 0.76 to 0.98 in triangular signals. Mean peak cross-correlations improved from 0.85 to 0.98 in random signals and from 0.93 to 0.98 for sinusoidal inputs, and mean time to peak cross-correlations decreased from 144 to 24 ms in random signals and from 156 to 25 ms in sine waves. Rise times in staircase signals decreased from a mean of 520 to 175 ms, and mean steady state error decreased from 12 to 3%. Significant effects of the triangle cycle time, sinusoidal frequency and staircase step were found on the performance of the muscle force control system. In addition, the possible effects of intrinsic feedback mechanisms on the control system were examined by repeating the closed loop part of the study but with the sciatic nerve cut proximally. The tracking results were essentially and statistically the same as in the closed loop condition. It was concluded that a simple feedback configuration provided superior tracking performance for a practical application in which orderly recruitment is used to control muscles; furthermore, it was concluded that this type of system would be virtually immune to external disturbances such as spasticity resulting from intact spinal neural feedback mechanisms found in paralyzed individuals.

Frequency domain-based models of skeletal muscle

Models of skeletal muscle based on its response to sinusoidal stimulation have been in use since the late 1960s. In these methods, cyclic excitation at varying frequencies is used to determine force or muscle length amplitude and phase as functions of excitation frequency. These functions can then be approximated by models consisting of combinations of poles and zeros and pure time delays without the need to combine force–length or force–velocity relationships. The major findings of a series of frequency response studies undertaken in our laboratory revealed that: • The frequency response models for isometric force including orderly recruitment of motor units were relatively invariant of the particular strategy or oscillation level employed. A critically damped second order model with corner frequency near 2 Hz and a pure time delay best described the relationship between input stimulation and output isometric force. • The frequency response models for load-moving muscles consisted of an overall gain which is a function of mass, dependent mostly on the width of the length–force relation at a given load (force), and a frequency-dependent gain component independent of load mass. The phase lag between input and output was also independent of load. • Muscle function and architecture are the primary determinants of its isometric force frequency response. • Tendon viscoelasticity (excluding the aponeurosis) has no significant effect on isometric force dynamic response, but does have a minor effect on load-moving dynamic response. The effect of tendon in reducing or augmenting the load-moving muscle response bandwidth is muscle-dependent. • The joint produces decreased high frequency gain and uniformly increased phase lags between input excitation and output force in isometric conditions. The joint acts as a lag network in load-moving conditions, increasing the phase lag without significant effect on the gain. Despite its inherent non-linear properties, the joint does not significantly deteriorate output signal quality in either isometric or load-moving conditions. • Co-contraction strategy has a significant effect on the dynamic response of the joint. These frequency-based models have shown to be robust as long as the excitation type and mechanical conditions under which they are obtained are not varied. They are particularly useful for the design of neuroprostheses, where a dynamic description of muscle output as a function of stimulus input under given conditions is desirable.

Muscle damage induced by stretch-shortening cycle exercise.

Strenuous stretch-shortening cycle exercise was used as a model to study the leakage of proteins from skeletal muscle. The analysis included serum levels of creatine kinase (S-CK), myoglobin (S-Mb), and carbonic anhydrase (S-CA III). Blood samples from power- (N=11) and endurance-trained (N=10) athletes were collected before, 0, and 2 h after the exercise, which consisted of a total of 400 jumps. The levels of all determined myocellular proteins increased immediately after the exercise (P < 0.05-0.001) among both subject groups. In the endurance group, the protein levels increased (P < 0.05-0.001) further during the following 2 h after the exercise, and the ratio of S-CA III and S-Mb decreased (P < 0.05) in a before-after comparison. This was not the case among the power group despite their greater mechanical work (P < 0.001) and higher ratio of eccentric and concentric EMG activity of the leg extensor muscles (P < 0.05). The differences of the determined protein levels between the subject groups might be due to obvious differences in the muscle fiber distribution, differences in recruitment order of motor units, and/or differences in training background.

Simulation of motor unit recruitment and microvascular unit perfusion: spatial considerations.

Muscle fiber activity is the principal stimulus for increasing capillary perfusion during exercise. The control elements of perfusion, i.e., microvascular units (MVUs), supply clusters of muscle fibers, whereas the control elements of contraction, i.e., motor units, are composed of fibers widely scattered throughout muscle. The purpose of this study was to examine how the discordant spatial domains of MVUs and motor units could influence the proportion of open capillaries (designated as perfusion) throughout a muscle cross section. A computer model simulated the locations of perfused MVUs in response to the activation of up to 100 motor units in a muscle with 40,000 fibers and a cross-sectional area of 100 mm2. The simulation increased contraction intensity by progressive recruitment of motor units. For each step of motor unit recruitment, the percentage of active fibers and the number of perfused MVUs were determined for several conditions: 1) motor unit fibers widely dispersed and motor unit territories randomly located (which approximates healthy human muscle), 2) regionalized motor unit territories, 3) reversed recruitment order of motor units, 4) densely clustered motor unit fibers, and 5) increased size but decreased number of motor units. The simulations indicated that the widespread dispersion of motor unit fibers facilitates complete capillary (MVU) perfusion of muscle at low levels of activity. The efficacy by which muscle fiber activity induced perfusion was reduced 7- to 14-fold under conditions that decreased the dispersion of active fibers, increased the size of motor units, or reversed the sequence of motor unit recruitment. Such conditions are similar to those that arise in neuromuscular disorders, with aging, or during electrical stimulation of muscle, respectively.

Motor unit recruitment order during voluntary and electrically induced contractions in the tibialis anterior.

The recruitment order of motor units (MU) was compared during voluntary and electrically induced contractions. With the use of spike-triggered averaging, a total of 302 MUs with recruitment thresholds ranging from 1% to 88% of maximal voluntary contraction were recorded in the human tibialis anterior muscle in five subjects. The mean (+/-SD) MU force was 98.3+/-93.3 mN (mean torque 16.8+/-15.9 mNm) and the mean contraction time (CT) 46.2+/-12.7 ms. The correlation coefficients (r) between MU twitch force and CT versus the recruitment threshold in voluntary contractions were +0.68 and -0.38 (P<0.001), respectively. In voluntary contractions, MUs were recruited in order of increasing size except for only 6% of the cases; whereas, during transcutaneous electrical stimulation (ES) at the muscle motor point, MU pairs showed a reversal of recruitment order in 28% and 35% of the observations, respectively, when the pulse durations were 1.0 ms or 0.1 ms. This recruitment reversal during ES was not related to the magnitude of the difference in voluntary recruitment thresholds between MUs. It is concluded that if the reversal of MU recruitment observed during ES is biophysically controlled by differences in their nerve axon input impedance, in percutaneous stimulation at the motor point, other factors such as the size and the morphological organisation of the axonal branches can also influence the order of activation.

Characteristic changes of motor unit activity in hip joint flexor muscles during voluntary isometric contraction during water immersion

The effect of whole body water immersion on the recruitment order of hip flexor motor units was investigated in 11 male subjects. Intramuscular spike potentials were recorded, with fine bipolar wire electrodes, from the iliopsoas, the sartorius, the rectus femoris and the tensor fasciae latae during voluntary isometric contraction while the subjects were standing erect with the hip on the test side flexed to 60 ° and the knee flexed to 120 °. Data were analysed by measuring the recruitment threshold in slow ramp contraction and by a computer-aided amplitude-frequency histogram of the spike potentials during short sustained contraction. The motor units were classified as low-amplitude units if they delivered spike potentials of less than 0.5 mV and high-amplitude units if the spike potentials exceeded 0.5 mV. In the ramp experiments, exposure to water immersion gave rise to a sudden increase in the recruitment thresholds of the low-amplitude units in all muscles, while in the recruitment thresholds of the high-amplitude units, the alterations differed among the muscles. The thresholds in the rectus femoris and tensor fasciae latae increased in the same direction as those of the low-amplitude units, while those in the iliopsoas and sartorius decreased in the opposite direction. The amplitude-frequency histograms clearly indicated that these different alterations occurred in all subjects, without exception. We concluded that unloading induced by water immersion changed the recruitment order of motor units during isometric contraction in the iliopsoas and sartorius, facilitating the recruitment of their larger motor units.

Activation of type-identified motor units during centrally evoked contractions in the cat medial gastrocnemius muscle. I. Motor-unit recruitment.

1. The recruitment order of 64 pairs of motor units, comprising 21 type-identified units, was studied during centrally evoked muscle contractions of the cat medial gastrocnemius (MG) muscle in an unanesthetized, high decerebrate preparation. Motor units were functionally isolated within the MG nerve by intra-axonal (or intramyelin) penetration with conventional glass microelectrodes. 2. Graded stimulation of the mesencephalic locomotor region (MLR) was used to evoke smoothly graded contractions, which under favorable conditions was estimated to reach 40% of maximum tetanic tension of the MG muscle. With this method of activation, 100% of slow twitch (type S) units, 95% of fast twitch, fatigue-resistant (type FR) units, 86% of fast twitch, fatigue-intermediate (type FI) units, and 49% of fast twitch, fatigable (type FF) units studied were recruited. 3. Motoneuron size as estimated by axonal conduction velocity (CV) was correlated with muscle-unit size as estimated by maximum tetanic tension (Po). Although the correlation between these properties was significant among type S and FR units, no significant correlation was found for these properties among type FI and FF units. 4. Motor-unit recruitment was ordered by physiological type (S > F, 100% of pairs; S > FR > FI > FF, 93% of pairs). Although none of the motor-unit properties studied predicted recruitment order perfectly, motor-unit recruitment was found to proceed by increasing Po (89% of pairs), decreasing contraction time (79% of pairs), decreasing fatigue index (80% of pairs), and increasing CV (76% of pairs). These percentages were significantly different from random (i.e., 50%). Statistically, all four motor-unit properties were equivalent in predicting recruitment order. These results are similar to those reported by other investigators for motor-unit recruitment order evoked from other supraspinal centers, as well as from peripheral sites. 5. When, however, motor-unit recruitment within pairs of motor units containing two fast-twitch (type F) units was examined, Po was a significantly better predictor of recruitment order than CV (85% vs. 52% of pairs). One explanation for this observation is that the correlation between Po and CV is high among type S, type FR units, and possibly among the lower-tension type FF units, but not among the remaining higher-tension type FF units. 6. The reproducibility of recruitment order in multiple contractions was investigated in 16 motor-unit pairs. Recruitment order was found to be variable in only three motor-unit pairs, all of which contained units of similar physiological type and recruitment threshold. 7. Analysis of recruitment order by pair-wise testing confirms the general conclusion reached in human studies that the muscle force level at recruitment for a motor unit is highly correlated with its strength. As an additional confirmation, the whole-muscle force level at recruitment for 41 units was measured in a series of contractions in which the rate of rise of muscle tension was limited to rates < 1,000 g/s. For these contractions, a significant correlation was found between muscle tension at recruitment and motor-unit Po.

Activation of type-identified motor units during centrally evoked contractions in the cat medial gastrocnemius muscle. II. Motoneuron firing-rate modulation.

1. The aim of this study was to examine the nature of motoneuron firing-rate modulation in type-identified motor units during smoothly graded contractions of the cat medial gastrocnemius (MG) muscle evoked by stimulation of the mesencephalic locomotor region (MLR). Motoneuron discharge patterns, firing rates, and the extent of firing-rate modulation in individual units were studied, as was the extent of concomitant changes in firing rates within pairs of simultaneously active units. 2. In 21 pairs of simultaneously active motor units, studied during 41 evoked contractions, the motoneurons' discharge rates and patterns were measured by processing the cells' recorded action potentials through windowing devices and storing their timing in computer memory. Once recruited, most motoneurons increased their firing rates over a limited range of increasing muscle tension and then maintained a fairly constant firing rate as muscle force continued to rise. Most motoneurons also decreased their firing rates over a slightly larger, but still limited, range of declining muscle force before they were derecruited. Although this was the most common discharge pattern recorded, several other interesting patterns were also seen. 3. The mean firing rate for slow twitch (type S) motor units (27.8 imp/s, 5,092 activations) was found to be significantly different from the mean firing rate for fast twitch (type F) motor units (48.4 imp/s, 11,272 activations; Student's t-test, P < 0.001). There was no significant difference between the mean firing rates of fast twitch, fatigue-resistant (type FR) and fast twitch, fatigable (type FF) motor units. When the relationship between motoneuron firing rate and whole-muscle force was analyzed, it was noted that, in general, smaller, lower threshold motor units began firing at lower rates and reached lower peak firing rates than did larger, higher threshold motor units. These results confirm both earlier experimental observations and predictions made by other investigators on the basis of computer simulations of the cat MG motor pool, but are in contrast to motor-unit discharge behavior recorded in some human motor-unit studies. 4. The extent of concomitant changes in firing rate within pairs of simultaneously active motor units was examined to estimate the extent of simultaneous motoneuron firing-rate modulation across the motoneuron pool. A smoothed (5 point sliding average) version of the two motoneurons' instantaneous firing rates was plotted against each other, and the slope and statistical significance of the relationship was determined. In 16 motor-unit pairs, the slope of the motoneurons' firing-rate relationship was significantly distinct from 0. Parallel firing-rate modulation (< 10-fold difference in firing rate change reflected by a slope of > 0.1) was noted only in pairs containing motor units of like physiological type and then only if they were of similar recruitment threshold. 5. Other investigators have demonstrated that changes in a motoneuron's "steady-state" firing rate predictably reflect changes in the amount of effective synaptic current that cell is receiving. The finding in the present study of limited parallel firing-rate modulation between simultaneously active motoneurons would suggest that changes in the synaptic drive to the various motoneurons of the pool is unevenly distributed. This finding, in addition to the findings of orderly motor-unit recruitment and the relationship between motor-unit recruitment threshold and motoneuron firing rate, cannot be adequately accommodated for by the existing models of the synaptic organization in motoneuron pools. Therefore a new model of the synaptic organization within the motoneuron pool has been proposed.