Muscle activity reduces soft-tissue resonance at heel-strike during walking

Muscle activity has previously been suggested to minimize soft-tissue resonance which occurs at heel-strike during walking and running. If this concept were true then the greatest vibration damping would occur when the input force was closest to the resonant frequency of the soft-tissues at heel-strike. However, this idea has not been tested. The purpose of this study was to test whether muscle activity in the lower extremity is used to damp soft-tissue resonance which occurs at heel-strike during walking. Hard and soft shoe conditions were tested in a randomized block design. Ground reaction forces, soft-tissue accelerations and myoelectric activity were measured during walking for 40 subjects. Soft-tissue mass was estimated from anthropologic measurements, allowing inertial forces in the soft-tissues to be calculated. The force transfer from the ground to the tissues was compared with changes in the muscle activity. The soft condition resulted in relative frequencies (input/tissue) to be closer to resonance for the main soft-tissue groups. However, no increase in force transmission was observed. Therefore, the vibration damping in the tissues must have increased. This increase concurred with increases in the muscle activity for the biceps femoris and lateral gastrocnemius. The evidence supports the proposal that muscle activity damps soft-tissue resonance at heel-strike. Muscles generate forces which act across the joints and, therefore, shoe design may be used to modify muscle activity and thus joint loading during walking and running.     

Modification of soft tissue vibrations in the leg by muscular activity.

Vibration characteristics were recorded for the soft tissues of the triceps surae, tibialis anterior, and quadriceps muscles. The frequency and damping of free vibrations in these tissues were measured while isometric and isotonic contractions of the leg were performed. Soft tissue vibration frequency and damping increased with both the force produced by and the shortening velocity of the underlying muscle. Both frequency and damping were greater in a direction normal to the skin surface than in a direction parallel to the major axis of each leg segment. Vibration characteristics further changed with the muscle length and between the individuals tested. The range of the measured vibration frequencies coincided with typical frequencies of impact forces during running. However, observations suggest that soft tissue vibrations are minimal during running. These results support the strategy that increases in muscular activity may be used by some individuals to move the frequency and damping characteristics of the soft tissues away from those of the impact force and thus minimize vibrations during walking and running.     

Tissue vibration in prolonged running

The impact force in heel–toe running initiates vibrations of soft-tissue compartments of the leg that are heavily dampened by muscle activity. This study investigated if the damping and frequency of these soft-tissue vibrations are affected by fatigue, which was categorized by the time into an exhaustive exercise. The hypotheses were tested that (H1) the vibration intensity of the triceps surae increases with increasing fatigue and (H2) the vibration frequency of the triceps surae decreases with increasing fatigue. Tissue vibrations of the triceps surae were measured with tri-axial accelerometers in 10 subjects during a run towards exhaustion. The frequency content was quantified with power spectra and wavelet analysis. Maxima of local vibration intensities were compared between the non-fatigued and fatigued states of all subjects. In axial (i.e. parallel to the tibia) and medio-lateral direction, most local maxima increased with fatigue (supporting the first hypothesis). In anterior–posterior direction no systematic changes were found. Vibration frequency was minimally affected by fatigue and frequency changes did not occur systematically, which requires the rejection of the second hypothesis. Relative to heel-strike, the maximum vibration intensity occurred significantly later in the fatigued condition in all three directions. With fatigue, the soft tissue of the triceps surae oscillated for an extended duration at increased vibration magnitudes, possibly due to the effects of fatigue on type II muscle fibers. Thus, the protective mechanism of muscle tuning seems to be reduced in a fatigued muscle and the risk of potential harm to the tissue may increase.     

Muscle activity damps the soft tissue resonance that occurs in response to pulsed and continuous vibrations.

This study tested the hypotheses that when the excitation frequency of mechanical stimuli to the foot was close to the natural frequency of the soft tissues of the lower extremity, the muscle activity increases 1) the natural frequency and 2) the damping to minimize resonance. Soft tissue vibrations were measured with triaxial accelerometers, and muscle activity was measured by using surface electromyography from the quadriceps, hamstrings, tibialis anterior, and triceps surae groups from 20 subjects. Subjects were presented vibrations while standing on a vibrating platform. Both continuous vibrations and pulsed bursts of vibrations were presented, across the frequency range of 10-65 Hz. Elevated muscle activity and increased damping of vibration power occurred when the frequency of the input was close to the natural frequency of each soft tissue. However, the natural frequency of the soft tissues did not change in a manner that correlated with the frequency of the input. It is suggested that soft tissue damping may be the mechanism by which resonance is minimized at heel strike during running.     

Changes in muscle activity in response to different impact forces affect soft tissue compartment mechanical properties

Electromyographic (EMG) activity is associated with several tasks prior to landing in walking and running including positioning the leg, developing joint stiffness and possibly control of soft tissue compartment vibrations. The concept of muscle tuning suggests one reason for changes in muscle activity pattern in response to small changes in impact conditions, if the frequency content of the impact is close to the natural frequency of the soft tissue compartments, is to minimize the magnitude of soft tissue compartment vibrations. The mechanical properties of the soft tissue compartments depend in part on muscle activations and thus it was hypothesized that changes in the muscle activation pattern associated with different impact conditions would result in a change in the acceleration transmissibility to the soft tissue compartments. A pendulum apparatus was used to systematically administer impacts to the heel of shod male participants. Wall reaction forces, EMG of selected leg muscles, soft tissue compartment and shoe heel cup accelerations were quantified for two different impact conditions. The transmissibility of the impact acceleration to the soft tissue compartments was determined for each subject/soft tissue compartment/shoe combination. For this controlled impact situation it was shown that changes in the damping properties of the soft tissue compartments were related to changes in the EMG intensity and/or mean frequency of related muscles in response to a change in the impact interface conditions. These results provide support for the muscle tuning idea--that one reason for the changes in muscle activity in response to small changes in the impact conditions may be to minimize vibrations of the soft tissue compartments that are initiated at heel-strike.     

Myoelectric signal conduction velocity and spectral parameters: influence of force and time

Reports on measurement of muscle fiber conduction velocity in humans are scarce. Inferences on the behavior of conduction velocity have been drawn from the behavior of myoelectric spectral parameters. The present report contains information on conduction velocity and spectral parameters studied at various muscle contraction levels and during and after sustained contractions. The following results have been obtained from measurements on the tibialis anterior muscle. Conduction velocity demonstrated a positive correlation with limb circumference and with muscle force output. Thus we conclude that the diameters of the muscle fibers of high-threshold motor units are, on an average, larger than those of low-threshold motor units. The study of a sustained contraction and on the recovery after such a contraction revealed that conduction velocity consistently decreased during a strong contraction as did various myoelectric spectral parameters. However, the spectral parameters decreased approximately twice as much as did the conduction velocity, and we conclude that factors other than the conduction velocity along the muscle fibers affect the myoelectric signal during a high-level contraction. These other factors appertain to changes in the firing statistics of individual motor units as well as the correlation between the firings of different motor units.     

Muscle activity in the leg is tuned in response to ground reaction forces.

During walking and running, the human body reacts to its external environment. One such response is to the impact forces that occur at heel strike. This study tested previous speculation that the levels of muscle activity in the lower extremities are adjusted in response to the loading rate of the impact forces. A pendulum apparatus was used to deliver repetitive impacts to the heels of 20 subjects. Impact forces were of similar magnitude to those experienced during running, but the loading rate was varied by 13% using different materials in the subjects' shoes. Myoelectric patterns were measured in the tibialis anterior, medial gastrocnemius, vastus medialis, and biceps femoris muscles. Wavelet analysis was used to resolve intensity of the myoelectric patterns into time and frequency space. Substantial and significant differences in the myoelectric activity occurred between the impact conditions for the 50 ms before and the 50 ms after impact, reaching 3 ms in timing, 16% in wavelet number, and 154% in the intensity of the muscle activity.     

Soft tissue vibrations within one soft tissue compartment

The concept of muscle tuning suggests that vibrations of the soft tissue compartments of the leg initiated by impacts are minimized by muscular activity prior to heel-strike of heel-toe running. For the quantification of muscle tuning it has been assumed (1) that the soft tissue compartment acts as one lumped mass and (2) that vibration energy dissipation does occur within one muscle. The purpose of this study was to test these two assumptions. It was hypothesized that (H1) the movement of the soft tissue compartment is not homogeneous, (H2) the vibration frequencies for different muscles within one soft tissue compartment are different and (3) attenuation of vibration movement within one muscle does occur. Soft tissue vibrations were measured using accelerometers on four locations on the quadriceps soft tissue compartment during heel-toe running. There were differences in the peak soft tissue acceleration and time of peak acceleration between accelerometer locations. The dominant frequency was similar throughout the soft tissue compartment, however; there was an attenuation of high-frequency vibration energy between distal and proximal points overlying one muscle. This evidence suggests that accelerometer placement is important when quantifying the acceleration magnitude and timing of peak soft tissue compartment but not when estimating the resonant vibration characteristics of a soft tissue compartment. It also provides initial evidence to support the idea that vibration control through muscle tuning may be achieved through changes in energy dissipating properties within the soft tissue compartment.     

Some mechanisms of voluntary control of walking in humans

The electrical activities of some muscles of the lower extremities have been studied by the method of presentation of an acoustic signal for a change in walking speed. It has been established that the motor response to a signal has two stages, at which (1) the ratio of muscle activities facilitating acceleration or deceleration of walking is formed and (2) the muscle activity corresponding to the new rate of locomotion is set. The latent period of the first stage of the motor response depends on the temporal relationship of the signal and the phase of muscle activity: it is minimum if the signal coincides with the phase of activity and maximum if the signal is given in the phase of bioelectric silence. It may be supposed that the voluntary control of the locomotion rate is related to at least two types of cortical effects: cyclic and acyclic. The former determine the transition from one speed of walking to another through changing human body posture characteristics (probably, they influence interneurons and motoneurons of reflex arcs); the latter, the characteristics of the new mode of the locomotor cycle by affecting the functional state of the interneurons and motoneurons of the spinal generator of stepping movements.     

Changes in EMG signals for the muscle tibialis anterior while running barefoot or with shoes resolved by non-linearly scaled wavelets

The purpose of this project was to study the EMG pattern of the tibialis anterior muscle in heel-toe running. Specifically, EMG changes in time, intensity and frequency shortly before and after heel-strike were addressed using an EMG-specific non-linearly scaled wavelets analysis. This method allowed extracting the time, intensity and frequency information inherent in the EMG signal at any time. The EMG signals of 40 male subjects were recorded for running barefoot and with shoes. The results confirmed that the pre-heel-strike EMG activities were typically seen at higher EMG frequencies (60-270Hz) while the post-heel-strike EMG activities resulted in lower frequency signals (10-90Hz). The timing of the pre-heel-strike EMG activities was not influenced by the used shoe conditions. The timing of the post-heel-strike EMG activities was significantly delayed when wearing shoes. The intensity of the pre-heel-strike muscle activity increased compared to the post-heel-strike one when wearing shoes. One can conclude that the activity of the tibialis anterior adjusts specifically to exterior conditions. The frequency shift between pre- and post heel-strike muscle activity were discussed with respect to activation of different motor units.     

Spectral properties of H-reflex recordings after an acute bout of whole-body vibration

Although research supports the use of whole-body vibration (WBV) to improve neuromuscular performance, the mechanisms for these improvements remain unclear. The purpose of this study was to identify the effect of WBV on the spectral properties of electrically evoked H-reflex recordings in the soleus (SOL) muscle. The H-reflex recordings were measured in the SOL muscle of 20 participants before and after a bout of WBV. The H-reflexes were evoked every 15 seconds for 150 seconds after WBV. A wavelet procedure was used to extract spectral data, which were then quantified with a principle components analysis. Resultant principle component scores were used for statistical analysis. The analysis extracted 1 principle component associated with the intensity of the myoelectric spectra and 1 principle component associated with the frequency. The scores of the principle component that were related to the myoelectric intensity were smaller at 30 and 60 milliseconds after WBV than before WBV. The WBV transiently decreased the intensity of myoelectric spectra during electrically evoked contractions, but it did not influence the frequency of the spectra. The decrease in intensity likely indicates a smaller electrically evoked muscle twitch response, whereas the lack of change in frequency would indicate a similar recruitment pattern of motor units before and after WBV.     

Interlimb interactions during cyclic in-phase and antiphase movements of arms and legs and their dependence on afferent influences

Coordinated arm and leg movements imply neural interactions between the rhythmic generators of the upper and lower extremities. In ten healthy subjects in the lying position, activity of the muscles of the upper and lower extremities was recorded during separate and joint cyclic movements of the arms and legs with different phase relationships between the movements of the limbs and under various conditions of the motor task. Antiphase active arm movements were characterized by higher muscle activity than during the inphase mode. The muscle activity during passive arm movements imposed by the experimentalist was significantly lower than muscle activity during passive arm movements imposed by the other arm. When loading one arm, the muscle activity in the other, passively moving, arm increased independently from the synergy of arm movements. During a motor task implementing joint antiphase movements of both upper and lower extremities, compared to a motor task implementing their joint in-phase movements, we observed a significant increase in activity in the biceps brahii muscle, the tibialis anterior muscle, and the biceps femoris muscle. Loading of arms in these motor tasks has been accompanied by increased activity in some leg muscles. An increase in the frequency of rhythmic movements resulted in a significant growth of the muscle activity of the arms and legs during their cooperative movements with a greater rate of rise in the flexor muscle activity of the arms and legs during joint antiphase movements. Thus, both the spatial organization of movements and the type of afferent influences are significant factors of interlimb interactions, which, in turn, determine the type of neural interconnections that are involved in movement regulation.     

Real-time gait assessment utilizing a new way of accelerometry

Real-time registration of body segment angles is essential in artificial body position control. A new method is presented for the real-time calculation of the lower extremity angles using data obtained from pairs of two one-dimensional accelerometers. It is shown that, assuming rigid-body dynamics and simple hinge joints, relative angles (i.e. angles between segments) can be calculated without integration, thereby solving the problem of integration drift normally associated with accelerometry. During the stance phase of walking, the relative angles can be transformed to absolute angles (i.e. relative to the gravitational field direction) for the different leg segments. The feasibility of relative angle calculation is demonstrated by calculation of the knee angle of a healthy subject. Stability and resolution were demonstrated with measurements during standing. Measurements during standing up, sitting down and walking showed that shock (heel-strike) and skin movements, due to movements of the underlying muscle tissue, are the main error sources. Additional signal processing, e.g. low-pass filtering, can be used to diminish this error. The accuracy of the knee angle found is shown to be high enough to be used in a feedback controller for functional electrostimulation of the lower extremities.     

The effect of activity during early postnatal development on motor unit size

At early stages of neuromuscular development, motor unit territory is expanded, with each muscle fibre being supplied by several axons. During postnatal development, some synapses are eliminated, motor unit size decreases, and the adult distribution of motor unit sizes emerges. This process depends on activity, since it proceeds more rapidly when the nerve is activated and is slower when activity is reduced. Here we studied whether, in addition to influencing the rate of retraction of motor unit territory, activity during the critical period of development affects the final outcome of the distribution of motor unit sizes. The sciatic nerve of 8- to 12-day-old rats was stimulated daily. One week later the tension of the extensor digitorum longus muscle and that of its individual motor units was recorded. The sizes of individual motor units were calculated and compared with those from animals that received no stimulation. The distribution of motor unit sizes from stimulated muscles was not significantly different from those from control muscles. Therefore, we conclude that although activity increases the rate at which motor units attain their adult size, it does not influence the final outcome of motor unit size distribution.     

Vibration settling time of the gastrocnemius remains constant during an exhaustive run in rear foot strike runners

In this study, vibrations of human gastrocnemius during an exhaustive run protocol are considered for analysis. Previous studies have shown increased vibration intensity and damping coefficient within the soft tissue with fatigue. The question of this study was to investigate if the vibration settling time remains constant during a prolonged running. Eleven semi-professional middle/long distance runners ran to exhaustion on a treadmill with their preferred constant speed (4.29 ± 0.33 m/s) for 3873 ± 1147 m. Vibration of the gastrocnemius lateralis, electrical activity of the tibialis anterior and the gastrocnemius medialis along with ground reaction force (GRF) were recorded. The results demonstrated significant increase in impact peak and loading rate, and the frequency content of the impact, with no significant change in active peak of the vertical GRF. Fatigue resulted in increased vibration intensity, damping coefficient, and energy dissipation of vibration with no change in vibration settling time. Furthermore, peak acceleration significantly linearly (R = 0.59) increased as a function of running time. The mean frequency of muscle activity of the gastrocnemius medialis and the intensity of muscle activity in TA significantly decreased. The results suggest that constant vibration settling time might either be an objective for muscle tuning which is more pronounced in fatigued state or a passive by-product of muscle function in running. Further studies are needed to address this point.     

Muscle fatigue - from motor units to clinical symptoms

Reductionist approaches have provided little insight on the fatigue experienced by humans during activities of daily living. Some of the reasons for this lack of progress include the persistence of outdated concepts, the misinterpretation of experimental recordings, and a failure to embrace a global perspective on fatigue. This paper summarizes the three examples of these limitations that were discussed in the 2011 Muybridge Award lecture: motor unit types and muscle fatigue, myoelectric manifestations of fatigue, and fatigue and fatigability. Although the motor units in a population do exhibit a range of fatigability values, there are not distinct groups of motor units and the concept that some motor units are resistant to fatigue emerged from protocols in which motor units were activated by electrical stimulation rather than voluntary activation. The concept of distinct motor unit types should be abandoned. The second example discussed in the lecture was the use of surface EMG signals to assess fatigue-related adjustments in motor unit activity. The critical assumption with this approach is that the association between surface EMG amplitude and muscle force remains constant during fatiguing contractions. Unfortunately, the relation does not remain constant and a series of computational studies demonstrate the magnitude of the discrepancy, including the absence of an association with the activation signal emerging from the spinal cord and that received by the muscle. The third example concerned the concepts of fatigue and fatigability. It has long been recognized that fatigue involves both sensations and impairments in motor function, and the final part of the lecture urged the integration of the two constructs into a single scheme in which fatigue can be modulated either independently or by interactions between perceptions of fatigue and the mechanisms that establish levels of fatigability. The expectation is that such critical evaluations of the concepts and approaches to the study of fatigue will provide a more effective foundation from which to identify the factors that contribute to fatigue in health and disease.     

Motor unit recruitment strategy of knee antagonist muscles in a step-wise, increasing isometric contraction

The purpose of this study was to determine if differences exist between the control strategies of two antagonist thigh muscles during knee flexion and extension muscular coactivation. Surface myoelectric signal (MES) of the quadriceps (rectus femoris) and the hamstrings (semitendinosus) were obtained from both muscles while performing step-wise increasing contractions during flexion and extension with the knee at 1.57 rad of flexion (90 degrees). The median frequency of the power density spectrum, which is related to the average muscle fiber action potential conduction velocity and therefore to motor unit recruitment, was calculated from each MES. The results suggest that, in all the subjects tested, when the muscle acts as antagonist most motor units are recruited up to 50% of the maximal voluntary force, whereas when the muscle acts as antagonist motor units are recruited up to 40% of the maximal voluntary force. The force range past 40–50% of the maximal force is also characterized by differences between the agonist/antagonist.     

Aging effects on the Achilles tendon moment arm during walking

The Achilles tendon (AT) moment arm transforms triceps surae muscle forces into a moment about the ankle which is critical for functional activities like walking. Moreover, the AT moment arm changes continuously during walking, as it depends on both ankle joint rotation and triceps surae muscle loading (presumably due to bulging of the muscle belly). Here, we posit that aging negatively effects the architecturally complex AT moment arm during walking, which thereby contributes to well-documented reductions in ankle moment generation during push-off. We used motion capture-guided ultrasound imaging to quantify instantaneous variations in the AT moment arms of young (23.9 ± 4.3 years) and older (69.9 ± 2.6 years) adults during walking, their dependence on triceps surae muscle loading, and their association with ankle moment generation during push-off. Older adults walked with 11% smaller AT moment arms and 11% smaller peak ankle moments during push-off than young adults. Moreover, as hypothesized, these unfavourable changes were significantly and positively correlated (r² = 0.38, p < 0.01). More surprisingly, aging attenuated load-dependent increases in the AT moment arm (i.e., those between heel-strike and push-off at the same ankle angle); only young adults exhibited a significant increase in their AT moment arm due to triceps surae muscle-loading. Age-associated reductions in triceps surae volume or activation, and thus muscle bulging during force generation, may compromise the mechanical advantage of the AT during the critical push-off phase of walking in older adults. Thus, strategies to restore and/or improve locomotor performance in our aging population should consider these functionally important changes in musculoskeletal behavior.     

Evidence of self-sustained motoneuron firing in young and older adults.

Motoneurons demonstrate a type of self-sustained firing behavior that seems to be produced by a prolonged period of depolarization caused by intrinsic long-term changes in the motoneuron. Such self-sustained firing behavior has previously been reported in human motor units. The purpose of the present study was to investigate the occurrence of self-sustained firing behavior in older adults. Eight young (mean age 24 yrs) and eight older (mean age 73 yrs) individuals participated in the investigation. While subjects produced light dorsiflexion contractions, a brief vibration stimulus was applied to the tibialis anterior muscle. Motor unit recordings were also obtained from the tibialis anterior muscle. Self-sustained firing behavior was evidenced by the appearance of new motor unit recruitment following vibration, even as the motor units that fired before the vibratory stimulus maintained a steady firing rate. The proportion of motor units exhibiting self-sustained firing activity was similar in both young and older adults (approx. 23% of trials). We conclude that self-sustained firing behavior is a ubiquitous phenomenon that does not seem to be affected by the aging process.     

Long-term use of high-heeled shoes alters the neuromechanics of human walking

Human movement requires an ongoing, finely tuned interaction between muscular and tendinous tissues, so changes in the properties of either tissue could have important functional consequences. One condition that alters the functional demands placed on lower limb muscle-tendon units is the use of high-heeled shoes (HH), which force the foot into a plantarflexed position. Long-term HH use has been found to shorten medial gastrocnemius muscle fascicles and increase Achilles tendon stiffness, but the consequences of these changes for locomotor muscle-tendon function are unknown. This study examined the effects of habitual HH use on the neuromechanical behavior of triceps surae muscles during walking. The study population consisted of 9 habitual high heel wearers who had worn shoes with a minimum heel height of 5 cm at least 40 h/wk for a minimum of 2 yr, and 10 control participants who habitually wore heels for less than 10 h/wk. Participants walked at a self-selected speed over level ground while ground reaction forces, ankle and knee joint kinematics, lower limb muscle activity, and gastrocnemius fascicle length data were acquired. In long-term HH wearers, walking in HH resulted in substantial increases in muscle fascicle strains and muscle activation during the stance phase compared with barefoot walking. The results suggest that long-term high heel use may compromise muscle efficiency in walking and are consistent with reports that HH wearers often experience discomfort and muscle fatigue. Long-term HH use may also increase the risk of strain injuries.