Quantitative EMG analysis to investigate synergistic coactivation of ankle and knee muscles during isokinetic ankle movement. Part 1: time amplitude analysis.

Synergy generally refers to the coordinated action of several motor elements to produce a specific motor task, either intentionally or automatically. One example is motor irradiation, a sudden spread of synergistic muscular coactivation resulting from a forceful single joint movement. To investigate this type of synergy pattern, a quantitative EMG approach was employed to characterize explicit neuromuscular synergy in the ankle-knee complex during maximal ankle isokinetic contraction. In the present study, isokinetic ankle contractions, both dorsiflexion and plantarflexion, at four different speeds (30, 60, 120, and 240 degrees/s) were studied in a normal adult population (N=11) to assess synergistic coactivation of the prime movers (tibialis anterior and gastrocnemius) and irradiated muscles (ipsilateral and contralateral rectus femoris and biceps femoris) of the ankle-knee complex. Electromyographic signals were collected with surface EMG electrodes and processed with traditional time-amplitude analysis to examine specific neural control strategies. The data generally supported several empirical assumptions common to neurological facilitation techniques. (1) Motor irradiation to the knee muscles due to ankle muscle isokinetic contraction was strongly directionally dependent. (2) Motor irradiation to the ipsilateral knee muscles due to ankle isokinetic contraction was speed dependent. (3) The prime movers demonstrated a similar control strategy, irrespective of different contraction speeds.     

EMG-angle relationship of the hamstring muscles during maximum knee flexion.

The aim of the present study was to investigate the EMG-joint angle relationship during voluntary contraction with maximum effort and the differences in activity among three hamstring muscles during knee flexion. Ten healthy subjects performed maximum voluntary isometric and isokinetic knee flexion. The isometric tests were performed for 5 s at knee angles of 60 and 90 degrees. The isokinetic test, which consisted of knee flexion from 0 to 120 degrees in the prone position, was performed at an angular velocity of 30 degrees /s (0.523 rad/s). The knee flexion torque was measured using a KIN-COM isokinetic dynamometer. The individual EMG activity of the hamstrings, i.e. the semitendinosus, semimembranosus, long head of the biceps femoris and short head of the biceps femoris muscles, was detected using a bipolar fine wire electrode. With isometric testing, the knee flexion torque at 60 degrees knee flexion was greater than that at 90 degrees. The mean peak isokinetic torque occurred from 15 to 30 degrees knee flexion angle and then the torque decreased as the knee angle increased (p<0.01). The EMG activity of the hamstring muscles varied with the change in knee flexion angle except for the short head of the biceps femoris muscle under isometric condition. With isometric contraction, the integrated EMGs of the semitendinosus and semimembranosus muscles at a knee flexion angle of 60 degrees were significantly lower than that at 90 degrees. During maximum isokinetic contraction, the integrated EMGs of the semitendinosus, semimembranosus and short head of the biceps femoris muscles increased significantly as the knee angle increased from 0 to 105 degrees of knee flexion (p<0.05). On the other hand, the integrated EMG of the long head of the biceps femoris muscle at a knee angle of 60 degrees was significantly greater than that at 90 degrees knee flexion with isometric testing (p<0.01). During maximum isokinetic contraction, the integrated EMG was the greatest at a knee angle between 15 and 30 degrees, and then significantly decreased as the knee angle increased from 30 to 120 degrees (p<0.01). These results demonstrate that the EMG activity of hamstring muscles during maximum isometric and isokinetic knee flexion varies with change in muscle length or joint angle, and that the activity of the long head of the biceps femoris muscle differs considerably from the other three heads of hamstrings.     

Properties of twitch motor units of the ankle extensor muscles in the bullfrog Rana catesbeiana

A study of the mechanical properties of the twitch motor units in the ankle extensor muscles of bullfrogs was undertaken to expand our view of the diversity of motor unit properties among vertebrates. Two muscles were chosen that represent a wide range of extensor function: the plantaris longus (PL) is a large muscle providing most of the force for ankle extension in hopping and swimming, and the tibialis posticus (TP) is relatively small and may act as an ankle stabilizer or be primarily postural in function. Both muscles have highly fatigable motor units, but also some (especially in TP) low or non-fatigable ones. Mean tetanic tensions of motor units in both muscles are relatively large as compared with those of mammals but are especially large in PL, No clear correlations were found between contraction times and either motor unit tetanic tensions or fatigability, nor did the motor units fall into clearly defined types based on any functional parameters. Overall contraction and relaxation times are slow compared with those of mammals and are somewhat slow compared to those of other frogs; unlike results from earlier studies, the large units of PL are slower than the small units of TP. This results in PL units reaching fused tetani at lower stimulus frequencies. The twitch/tetanus and force/frequency ratios in PL motor units are much larger than those of TP, giving PL units greater relative forces at lower stimulus rates. These results are discussed in the context of motor unit function. © 1994 Wiley-Liss, Inc.     

Effect of fatigue on hamstring coactivation during isokinetic knee extensions

We examined the effect of fatigue of the quadriceps muscles on coactivation of the hamstring muscles and determined if the response is different between two isokinetic speeds in ten males and ten females with no history of knee pathology. Electromyographic data were recorded from the vastus lateralis and biceps femoris muscles during 50 maximal knee extensions at isokinetic speeds of 1.75 rad · s⁻¹ (100° · s⁻¹) and 4.36 rad · s⁻¹ (250° · s⁻¹). A greater degree of coactivation was apparent at the higher speed, but the increase in coactivation of the hamstring muscles was similar at both speeds. The results revealed that: (1) coactivation is greater at a higher isokinetic speed, and (2) coactivation increases during fatigue, but the rate of increase is independent of contraction velocity. Accepted: 15 June 1998     

Effect of contraction form and contraction velocity on the differences between resultant and measured ankle joint moments

During a maximal isometric plantar flexion effort the moment measured at the dynamometer differs from the resultant ankle joint moment. The present study investigated the effects of contraction form and contraction velocity during isokinetic plantar/dorsal flexion efforts on the differences between resultant and measured moments due to the misalignment between ankle and dynamometer axes. Eleven male subjects (age: 31+/-6 years, mass: 80.6+/-9.6 kg, height: 178.4+/-7.4 cm) participated in this study. All subjects performed isometric-shortening-stretch-isometric contractions induced by electrical stimulation at three different angular velocities (25 degrees /s, 50 degrees /s and 100 degrees /s) on a customised dynamometer. The kinematics of the leg were recorded using the vicon 624 system with eight cameras operating at 250 Hz. The resultant moments at the ankle joint were calculated through inverse dynamics. The relative differences between resultant and measured ankle joint moments due to axis misalignment were fairly similar in all phases of the isometric-shortening-stretch-isometric contraction (in average 5-9% of the measured moment). Furthermore these findings were independent of the contraction velocity. During dynamic plantar/dorsal flexion contractions the differences between measured and resultant joint moment are high enough to influence conclusions regarding the mechanical response of ankle extensor muscles. However the relative differences were not increased during dynamic contractions as compared to isometric contractions.     

Effect of hip and ankle muscle fatigue on unipedal postural control

Fatigue and deficits in postural control may predispose musculoskeletal injury. The purpose of this study was to examine the effects of fatigue at the hip and ankle during frontal plane movements on postural control during single-leg stance. Thirteen healthy volunteers completed two testing sessions 1 week apart consisting of isokinetic fatigue of the frontal plane movers of either the ankle or hip with measures of static unipedal postural control taken before and after fatigue. Postural control was assessed during three 30-s trials of unilateral stance with eyes open before and after the fatigue protocol at each testing session. Mean center of pressure (COP) excursion velocity in the sagittal and frontal planes was compared between pre- and post-fatigue across the two joints. Fatigue of the hip musculature led to postural control impairments in the frontal and sagittal planes, while fatigue of the ankle musculature did not significantly impair postural control in either plane. Our results suggest that there is a greater effect of localized fatigue of the frontal plane movers of the hip compared to the ankle on maintenance of a postural control in single-leg stance.     

Electromyogram power spectra frequencies associated with motor unit recruitment strategies

The isolated contributions of motor unit recruitment and firing rate variations to the median frequency of the electromyogram's power density spectrum were determined. Orderly stimulation of the cat gastrocnemius motor units via nerve electrodes gave rise to linearly increasing median frequency regardless of the action potential firing rate of the active motor units. Increase in the discharge rate of all the motor units resulted in nearly constant median frequency. It was concluded that the increasing average conduction velocity during motor unit recruitment is the major contributor to variations in the electromyogram median frequency. The possibility of using the median frequency as the index to identify the recruitment control strategies employed by various muscles during increasing force contraction is suggested.     

Human group I excitatory projections from ankle dorsiflexors to quadriceps muscle

A short latency projection of group I afferent fibers from ankle dorsiflexors to knee extensor muscles has been categorized as species specific to humans. However, the effects of the pathway have only been inferred from conditioning homonymous reflexes in relaxed muscle. This study focused directly on the responses evoked in the electromyogram of the heteronymous muscles when active, in two experiments. In the first, preferential activation of group I afferents of ankle dorsiflexors, by electrical stimulation of the common peroneal nerve, excited both vastus medialis (mean latency, 26.3 ms) and rectus femoris (mean latency, 33.5 ms). No excitation or inhibition in either muscle was associated with stimulation of the tibial nerve. The second experiment compared vastus medialis responses with common peroneal nerve stimulation during three different movement conditions in which the muscle was equally contracted: rhythmic isotonic (pedalling); episodic isotonic; isometric contraction. Responses were identified in all three active states, with no significant differences in amplitude or latency. No responses were seen in the relaxed muscle.     

Electromyographic normalization procedures for determining exercise intensity of closed chain exercises for strengthening the quadriceps femoris muscles

The purpose of this study was to compare the electromyographic (EMG) amplitudes of the quadriceps femoris (QF) muscles during a maximum voluntary isometric contraction (MVIC) to submaximal and maximal dynamic concentric contractions during active exercises. A secondary purpose was to provide information about the type of contraction that may be most appropriate for normalization of EMG data if one wants to determine if a lower extremity closed chain exercise is of sufficient intensity to produce a strengthening response for the QF muscles. Sixty-eight young healthy volunteers (39 female, 29 male) with no lower extremity pain or injury participated in the study. Surface electrodes recorded EMG amplitudes from the vastus medialis obliquus (VMO), rectus femoris (RF), and vastus lateralis (VL) muscles during 5 different isometric and dynamic concentric exercises. The last 27 subjects performed an additional 4 exercises from which a second data set could be analyzed. Maximum isokinetic knee extension and moderate to maximum closed chain exercises activated the QF significantly more than a MVIC. A 40-cm. lateral step-up exercise produced EMG amplitudes of the QF muscles of similar magnitude as the maximum isokinetic knee extension exercises and would be an exercise that could be considered for strengthening the QF muscles. Most published EMG studies of exercises for the QF have been performed by comparing EMG amplitudes during dynamic exercises to a MVIC. This procedure can lead one to overestimate the value of a dynamic exercise for strengthening the QF muscles. We suggest that when studying the efficacy of a dynamic closed chain exercise for strengthening the QF muscles, the exercise be normalized to a dynamic maximum muscle contraction such as that obtained with knee extension during isokinetic testing.     

The nature of facilitation of leg muscle motor evoked potentials by knee flexion

Knee flexion is a movement that initiates rising from a sitting position, which is a common therapeutic exercise for patients unable to ambulate. We investigated how voluntary isometric biceps femoris contraction affects motor evoked potential (MEP) amplitude following transcranial magnetic stimulation, background electromyographic (EMG) amplitude, and H-reflex amplitude in ipsilateral leg muscles. Subjects were seated on the edge of a bed with their hips and knees flexed at 90°, and the soles of their feet on the floor. MEP and background EMG were recorded from the tibialis anterior (TA) and soleus (SOL), and H reflexes from SOL of 30 volunteers. Background EMG and MEP also were recorded while voluntarily contracting tested muscles. Biceps femoris contraction increased MEP and background EMG for TA and SOL ( p < 0.01). Maximal background EMG and MEP increased with increasing voluntary contraction of tested muscles ( p < 0.005). Regression slope differed little between TA and SOL. Biceps femoris contraction facilitated MEP comparably for TA and SOL, while SOL background EMG exceeded that of TA ( p < 0.02). The relationship between MEP facilitation and background EMG changed to favor more efficient facilitation in TA ( p < 0.05), but not SOL ( p > 0.1). MEP recorded from TA and SOL with subthreshold stimuli using needle electrodes were more frequent with biceps femoris contraction ( p < 0.04). H-reflex amplitude of SOL decreased during biceps femoris contraction ( p < 0.001). We concluded that biceps femoris contraction affects leg muscle MEP, background EMG, and H reflexes differently.     

The effects of sloped surfaces on locomotion: an electromyographic analysis

Investigations using quadrupeds have suggested that the motor programs used for slope walking differ from that used for level walking. This idea has not yet been explored in humans. The aim of this study was to use electromyographic (EMG) signals obtained during level and slope walking to complement previously published joint angle and joint moment data in elucidating such control strategies. Nine healthy volunteers walked on an instrumented ramp at each of five grades (-39%, -15%, 0%, +15%, +39%). EMG activity was recorded unilaterally from eight lower limb muscles (gluteus maximus (GM), rectus femoris (RF), vastus medialis (VM), biceps femoris (BF), semimembranosus (SM), soleus (Sol), medial gastrocnemius (MG), and tibialis anterior (TA)). The burst onset, duration, and mean activity were calculated for each burst in every trial. The burst characteristics were then averaged within each grade and subject and submitted to repeated measures ANOVAs to assess the effect of grade (alpha=0.05, a priori). Power production increased during upslope walking, as did the mean activity and burst durations of most muscles. In this case, the changes in muscle activity patterns were not predictable based on the changes in joint moments because of the activation of biarticular muscles as antagonists. During downslope walking power absorption increased, as did knee extensor activity (mean and duration) and the duration of the ankle plantarflexor activity. The changes in muscle activity during this task were directly related to the changes in joint moments. Collectively these data suggest that the nervous system uses different control strategies to successfully locomote on slopes, and that joint power requirements are an important factor in determining these control strategies.     

Homologous Muscle Contraction during Unilateral Movement Does Not Show a Dominant Effect on Leg Representation of the Ipsilateral Primary Motor Cortex

Co-activation of homo- and heterotopic representations in the primary motor cortex (M1) ipsilateral to a unilateral motor task has been observed in neuroimaging studies. Further analysis showed that the ipsilateral M1 is involved in motor execution along with the contralateral M1 in humans. Additionally, transcranial magnetic stimulation (TMS) studies have revealed that the size of the co-activation in the ipsilateral M1 has a muscle-dominant effect in the upper limbs, with a prominent decline of inhibition within the ipsilateral M1 occurring when a homologous muscle contracts. However, the homologous muscle-dominant effect in the ipsilateral M1 is less clear in the lower limbs. The present study investigates the response of corticospinal output and intracortical inhibition in the leg representation of the ipsilateral M1 during a unilateral motor task, with homo- or heterogeneous muscles. We assessed functional changes within the ipsilateral M1 and in corticospinal outputs associated with different contracting muscles in 15 right-handed healthy subjects. Motor tasks were performed with the right-side limb, including movements of the upper and lower limbs. TMS paradigms were measured, consisting of short-interval intracortical inhibition (SICI) and recruitment curves (RCs) of motor evoked potentials (MEPs) in the right M1, and responses were recorded from the left rectus femoris (RF) and left tibialis anterior (TA) muscles. TMS results showed that significant declines in SICI and prominent increases in MEPs of the left TA and left RF during unilateral movements. Cortical activations were associated with the muscles contracting during the movements. The present data demonstrate that activation of the ipsilateral M1 on leg representation could be increased during unilateral movement. However, no homologous muscle-dominant effect was evident in the leg muscles. The results may reflect that functional coupling of bilateral leg muscles is a reciprocal movement.     

Spinal reflexes and coactivation of ankle muscles during a submaximal fatiguing contraction.

This study examined the involvement of spinal mechanisms in the control of coactivation during a sustained contraction of the ankle dorsiflexors at 50% of maximal voluntary contraction. Changes in the surface electromyogram (EMG) of the tibialis anterior and of two antagonist muscles, the soleus and lateral gastrocnemius, were investigated during and after the fatigue task. Concurrently, the compound action potential (M-wave) and the Hoffmann reflex of the soleus and lateral gastrocnemius were recorded. The results showed that the torque of the ankle dorsiflexors and the average EMG of the tibialis anterior during maximal voluntary contraction declined by 40.9 +/- 17.7% (mean +/- SD; P < 0.01) and 37.0 +/- 19.9% (P < 0.01), respectively, at task failure. During the submaximal fatiguing contraction, the average EMG of both the agonist and antagonist muscles increased, leading to a nearly constant ratio at the end of the contraction when normalized to postfatigue values. In contrast to the monotonic increase in average EMG of the antagonist muscles, the excitability of their spinal reflex pathways exhibited a biphasic modulation. The amplitude of the Hoffman reflexes in the soleus and lateral gastrocnemius increased to 147.5 +/- 52.9% (P < 0.05) and 166.7 +/- 74.9% (P < 0.01), respectively, during the first 20% of the contraction and then subsequently declined to 66.3 +/- 44.8 and 74.4 +/- 44.2% of their initial values. In conclusion, the results show that antagonist coactivation did not contribute to task failure. The different changes in voluntary EMG activity and spinal reflex excitability in the antagonist muscles during the fatiguing contraction support the concept that the level of coactivation is controlled by supraspinal rather than spinal mechanisms. The findings indicate, however, that antagonist coactivation cannot simply be mediated by a central descending "common drive" to the motor neuron pools of the agonist-antagonist muscle pairs. Rather, they suggest a more subtle regulation of the drive, possibly through presynaptic mechanisms, to the motoneurons that innervate the antagonist muscles.     

Mechanomyogram from the different heads of the quadriceps muscle during incremental knee extension

To investigate the time- and frequency-domain responses of mechanomyograms (MMGs) during the progressive fatigue induced by intermittent incremental contractions, a surface MMG was obtained from the three muscle heads of the quadriceps muscle in seven subjects while they performed isometric knee extensions lasting 7.6 min. Isometric intermittent incremental contractions started at 1% of the maximal voluntary contraction (MVC) for 3 s, with a 3-s relaxation period in between each contraction, and the contraction level was increased by 1% of MVC for every contraction (by 10% of MVC per min) up to exhaustion. Separate contractions with sufficient rest periods were also conducted to serve for the MMG characteristics without fatigue. The integrated MMG (iMMG) was linearly related to force in all of the muscles when fatigue was not involved. With regard to the incremental contractions, the relationship exhibited an ascending-descending shape, but the behavior was not the same for the individual muscle heads, especially for the rectus femoris muscle. A steep increase in the median frequency of MMG from around 60% of MVC corresponded to a decrease in iMMG. These results suggest that analysis of MMG in the time- and frequency-domain during an incremental protocol is a useful way of characterizing the motor unit recruitment strategy and fatigue properties of individual muscles. Accepted: 19 March 1998     

Motor unit firing patterns of lower leg muscles during isometric plantar flexion with flexed knee joint position

The ankle flexor and extensor muscles are essential for pedal movements associated with car driving. Neuromuscular activation of lower leg muscles is influenced by the posture during a given task, such as the flexed knee joint angle during car driving. This study aimed to investigate the influence of flexion of the knee joint on recruitment threshold-dependent motor unit activity in lower leg muscles during isometric contraction. Twenty healthy participants performed plantar flexor and dorsiflexor isometric ramp contractions at 30 % of the maximal voluntary contraction (MVC) with extended (0°) and flexed (130°) knee joint angles. High-density surface electromyograms were recorded from medial gastrocnemius (MG), soleus (SOL), and tibialis anterior (TA) muscles and decomposed to extract individual motor units. The torque-dependent change (Δpps /Δ%MVC) of the motor unit activity of MG (recruited at 15 %MVC) and SOL (recruited at 5 %MVC) muscles was higher with a flexed compared with an extended knee joint (p < 0.05). The torque-dependent change of TA MU did not different between the knee joint angles. The motor units within certain limited recruitment thresholds recruited to exert plantar flexion torque can be excited to compensate for the loss of MG muscle torque output with a flexed knee joint.     

Acute changes in motor unit discharge property after concentric versus eccentric contraction exercise in knee extensor

This study aimed to investigate the motor unit firing property immediately after concentric or eccentric contraction exercise. Eighteen healthy men performed repetitive maximal isokinetic knee extension exercises with only concentric or eccentric contraction until they exerted less than 80% of the baseline strength. Before and after the fatiguing exercise, high-density surface electromyography of the vastus lateralis was recorded during submaximal ramp-up isometric contraction and individual motor units were identified. Only motor units that could be tracked before and after exercise were analyzed. Muscle cross-sectional area of the vastus lateralis was measured using ultrasound, and electrically evoked torque was recorded before and after the exercise. Sixty-five and fifty-three motor units were analyzed before and after the concentric and eccentric contractions, respectively. The results showed that motor units with moderate to high recruitment thresholds significantly decreased recruitment thresholds under both conditions, and the motor unit discharge rates significantly increased after concentric contraction compared to eccentric contraction. A greater muscle cross-sectional area was observed with concentric contraction. The evoked torque was significantly decreased under both conditions, but no difference between the conditions. These results suggest that fatiguing exercise with concentric contraction contributes to greater neural input to muscles and metabolic responses than eccentric contraction.     

Triceps surae short latency stretch reflexes contribute to ankle stiffness regulation during human running

During human running, short latency stretch reflexes (SLRs) are elicited in the triceps surae muscles, but the function of these responses is still a matter of controversy. As the SLR is primarily mediated by Ia afferent nerve fibres, various methods have been used to examine SLR function by selectively blocking the Ia pathway in seated, standing and walking paradigms, but stretch reflex function has not been examined in detail during running. The purpose of this study was to examine triceps surae SLR function at different running speeds using Achilles tendon vibration to modify SLR size. Ten healthy participants ran on an instrumented treadmill at speeds between 7 and 15 km/h under 2 Achilles tendon vibration conditions: no vibration and 90 Hz vibration. Surface EMG from the triceps surae and tibialis anterior muscles, and 3D lower limb kinematics and ground reaction forces were simultaneously collected. In response to vibration, the SLR was depressed in the triceps surae muscles at all speeds. This coincided with short-lasting yielding at the ankle joint at speeds between 7 and 12 km/h, suggesting that the SLR contributes to muscle stiffness regulation by minimising ankle yielding during the early contact phase of running. Furthermore, at the fastest speed of 15 km/h, the SLR was still depressed by vibration in all muscles but yielding was no longer evident. This finding suggests that the SLR has greater functional importance at slow to intermediate running speeds than at faster speeds.     

Application of surface electromyography in assessing muscle recruitment patterns in a six-minute continuous rowing effort

Specific sequences of muscle coordination exist in movements of every sport. In particular, sports involving repetitive movement patterns such as rowing may rely more heavily on coordinated muscle contraction sequencing in order to produce optimal performance. The aim of this study was to monitor the fatigue patterns of the major muscles engaged during the rowing stroke in rowers of varying abilities during a 6-minute continuous maximal rowing effort on a Concept II rowing ergometer. Sixteen male rowers were categorized into 5 groups based on years of training and their average pace of the 6-minute continuous maximal rowing effort. Continuous surface electromyography signals, recorded from brachioradialis, biceps brachii, middeltoid, rectus abdominis, erector spinae, rectus femoris, biceps femoris, and medial gastrocnemius, were used to investigate the influence of local muscle fatigue on optimal muscle coordination sequences during the rowing exercise. Rowers who performed better on the ergometer test and had more rowing experience tended to portray muscle recruitment patterning, which alternately emphasized different major muscle groups in a form of sharing of workload. This sharing allowed mean peak frequency restitution to take place in some muscles, while others took on more of the workload. The muscles of rowers with less experience and lower levels of performance did not appear to exhibit this same phenomenon known as biodynamic compensation. If coaches have a clearer picture of the fatigue patterns and recruitment strategies occurring in their athletes during a maximal effort row, strength training program adaptations could be made to compensate for weaker areas, which may assist rowers in attaining and sustaining more optimal patterns and strategies throughout the exercise effort.     

Crossed inhibition as revealed by cross-correlogram between bilateral homonymous motor unit spikes in man.

Neuromuscular unit (NMU) spikes activated during the tonic vibration reflex (TVR) in man were studied by means of the cross-correlogram test. Those NMUs which preferredly fired at some definite interval with good correlation with vibration were called "locked" spikes. The amount above zero correlation level seen in the cross-correlogram was expressed as ipsilateral "excitatory frequency", Ef. TVR contraction of the ipsilateral quadriceps femoris muscle was stimulated by voluntary contraction of the contralateral quadriceps femoris. I. Cross-correlogram test of the contralateral voluntary NMU spikes with ipsilateral vibration revealed "contralateral inhibitory modulation". Such inhibitory modulation was observed in 24% of the contralateral NMUs during voluntary tracking contraction with visual feed-back and in 22% of NMUs during blind simulated contraction without visual feed-back. The "subtracted" amount, i.e., contralateral inhibitory modulation, was also expressed as the contralateral "inhibitory frequency", If, which was calculated from the indentations of the cross-correlogram. 2. When varied vibratory frequency was applied to the muscle both Ef and If attained their maximum value around an "optimal frequency" of vibration, i.e., 80-95 HZ. 3. The ratio Ef/If was proposed as the reciprocal co-activation ratio which was fairly independent of the change of vibratory frequency. An average value of Ef/If ratio was calculated at 2.2 +/- 0.9 from 19 experiments.