Symphyseal fusion and jaw-adductor muscle force: an EMG study

The purpose of this study is to test various hypotheses about balancing-side jaw muscle recruitment patterns during mastication, with a major focus on testing the hypothesis that symphyseal fusion in anthropoids is due mainly to vertically- and/or transversely-directed jaw muscle forces. Furthermore, as the balancing-side deep masseter has been shown to play an important role in wishboning of the macaque mandibular symphysis, we test the hypothesis that primates possessing a highly mobile mandibular symphysis do not exhibit the balancing-side deep masseter firing pattern that causes wishboning of the anthropoid mandible. Finally, we also test the hypothesis that balancing-side muscle recruitment patterns are importantly related to allometric constraints associated with the evolution of increasing body size. Electromyographic (EMG) activity of the left and right superficial and deep masseters were recorded and analyzed in baboons, macaques, owl monkeys, and thick-tailed galagos. The masseter was chosen for analysis because in the frontal projection its superficial portion exerts force primarily in the vertical (dorsoventral) direction, whereas its deep portion has a relatively larger component of force in the transverse direction. The symphyseal fusion-muscle recruitment hypothesis predicts that unlike anthropoids, galagos develop bite force with relatively little contribution from their balancing-side jaw muscles. Thus, compared to galagos, anthropoids recruit a larger percentage of force from their balancing-side muscles. If true, this means that during forceful mastication, galagos should have working-side/balancing-side (W/B) EMG ratios that are relatively large, whereas anthropoids should have W/B ratios that are relatively small. The EMG data indicate that galagos do indeed have the largest average W/B ratios for both the superficial and deep masseters (2.2 and 4.4, respectively). Among the anthropoids, the average W/B ratios for the superficial and deep masseters are 1.9 and 1.0 for baboons, 1.4 and 1.0 for macaques, and both values are 1.4 for owl monkeys. Of these ratios, however, the only significant difference between thick-tailed galagos and anthropoids are those associated with the deep masseter. Furthermore, the analysis of masseter firing patterns indicates that whereas baboons, macaques and owl monkeys exhibit the deep masseter firing pattern associated with wishboning of the macaque mandibular symphysis, galagos do not exhibit this firing pattern. The allometric constraint-muscle recruitment hypothesis predicts that larger primates must recruit relatively larger amounts of balancing-side muscle force so as to develop equivalent amounts of bite force. Operationally this means that during forceful mastication, the W/B EMG ratios for the superficial and deep masseters should be negatively correlated with body size. Our analysis clearly refutes this hypothesis. As already noted, the average W/B ratios for both the superficial and deep masseter are largest in thick-tailed galagos, and not, as predicted by the allometric constraint hypothesis, in owl monkeys, an anthropoid whose body size is smaller than that of thick-tailed galagos. Our analysis also indicates that owl monkeys have W/B ratios that are small and more similar to those of the much larger-sized baboons and macaques. Thus, both the analysis of the W/B EMG ratios and the muscle firing pattern data support the hypothesis that symphyseal fusion and transversely-directed muscle force in anthropoids are functionally linked. This in turn supports the hypothesis that the evolution of symphyseal fusion in anthropoids is an adaptation to strengthen the symphysis so as to counter increased wishboning stress during forceful unilateral mastication. (ABSTRACT TRUNCATED)     

Towards optimal multi-channel EMG electrode configurations in muscle force estimation: a high density EMG study.

Surface EMG is an important tool in biomechanics, kinesiology and neurophysiology. In neurophysiology the concept of high-density EMG (HD-EMG), using two dimensional electrode grids, was developed for the measurement of spatiotemporal activation patterns of the underlying muscle and its motor units (MU). The aim of this paper was to determine, with the aid of a HD-EMG grid, the relative importance of a number of electrode sensor configurations for optimizing muscle force estimation. Sensor configurations are distinguished in two categories. The first category concerns dimensions: the size of a single electrode and the inter electrode distance (IED). The second category concerns the sensor's spatial distribution: the total area from which signals are obtained (collection surface) and the number of electrodes per cm(2) (collection density). Eleven subjects performed isometric arm extensions at three elbow angles and three contraction levels. Surface-EMG from the triceps brachii muscle and the external force at the wrist were measured. Compared to a single conventional bipolar electrode pair, the force estimation quality improved by about 30% when using HD-EMG. Among the sensor configurations, the collection surface alone appeared to be responsible for the major part of the EMG based force estimation quality by improving it with 25%.     

Muscle force recruitment and biomechanical modeling: an analysis of masseter muscle function during mastication in Macaca fascicularis.

The main purpose of this study is to test the hypothesis that as subjects chew with increasing levels of force, the ratio of the working- to balancing-side jaw-muscle force (W/B) decreases and begins to approach 1.0. We did this by analyzing relative masseter force in Macaca fascicularis using both strain gage and surface electromyographic (EMG) techniques. In addition, we also analyzed: 1) the relationship between jaw position using cineradiographic techniques and relative masseter force, 2) the timing differences between relative masseter force from the working and balancing sides, and 3) the loading and unloading characteristics of the masseter muscle. Our findings indicate that when macaques increase the amount of overall masticatory force during chewing, the W/B ratio for masseter force frequently (but not always) decreases and begins to approach 1.0. Therefore, our working hypothesis is not completely supported because the W/B ratio does not decrease with increasing levels of force in all subjects. The data also demonstrate timing differences in masseter force. During apple-skin mastication, the average peak masseter force on the working side occurs immediately at or slightly after the initial occurrence of maximum intercuspation, whereas the average peak masseter force on the balancing side occurs well before maximum intercuspation. On average, we found that peak force from the balancing-side masseter precedes the working-side masseter by about 26 msec. The greater the asynchrony between working- and balancing-side masseter force, the greater the difference in the relative magnitude of these forces. For example, in the subject with the greatest asynchrony, the balancing-side masseter had already fallen to about one-half of peak force when the working-side masseter reached peak force. Our data also indicate that the loading and unloading characteristics of the masseter differ between the working and balancing sides. Loading (from 50 to 100% of peak force) and unloading (from 100 to 50% of peak force) for the balancing-side masseter tends to be rather symmetrical. In contrast, the working-side masseter takes much longer to load from 50 to 100% of peak force than it does to unload from 100 to 50% of peak force. Finally, it takes on average about 35 msec for the working-side zygoma and 42 msec for the balancing-side zygoma to unload from 100 to 50% of peak force during apple-skin mastication, indicating that the unloading characteristics of the macaque masseter during mastication closely approximates its relaxation characteristics (as determined by muscle stimulation).     

Testing of a biomechanical model of the lumbar muscle force distribution using quasi-static loading exercises.

The study of lumbar muscle force distribution in response to externally applied loads is based on the introduction of biomechanical models of the lumbar region. The evaluation of such models requires the execution of loading exercises while monitoring the EMG activity of certain lumbar muscles. This work uses muscle activity maps as the major design tool of such exercises, provided that the subject is constrained to an upright erect posture. The maps describe the predicted muscle force for a given combination of externally applied bending moments. A series of shoulder adduction exercises were designed and the EMG signals of eight lumbar muscles were measured while subjects performed the exercises. The results show good agreement between the model predictions and the EMG measurements, especially when the load and the muscle were contralateral to one another.     

Heel-off perturbation during gait initiation: biomechanical analysis using triaxial accelerometry and a force plate.

This study analyzes the movements of the hips, shoulders and of the body center of gravity before and at heel-off, when step execution begins to initiate gait from an upright posture. The heel-off movement was considered as a dynamic perturbation induced by the stepping movement. The experimental paradigm used for studying this perturbation was the single-step movement, in which the initial posture and voluntary movements are identical to those of gait initiation. Data were collected from accelerometer recordings of the triaxial accelerations at the joints of the upper part of the body, and by calculating the triaxial accelerations of the center of gravity using force plate measurements. The resultant vectors were used to establish and compare the magnitude and direction of the accelerations at different joints, and from them, the roles of the pelvis and the scapular girdles with respect to the objectives of the gait movement.     

Interpreting muscle function from EMG: lessons learned from direct measurements of muscle force

Electromyography is often used to infer the pattern of productionof force by skeletal muscles. The interpretation of muscle functionfrom the electromyogram (EMG) is challenged by the fact thatfactors such as type of muscle fiber, muscle length, and musclevelocity can all influence the relationship between electricaland mechanical activity of a muscle. Simultaneous measurementsof EMG, muscle force, and fascicle length in hindlimb musclesof wild turkeys allow us to probe the quantitative link betweenforce and EMG. We examined two features of the force–EMGrelationship. First, we measured the relaxation electromechanicaldelay (r-EMD) as the time from the end of the EMG signal totime of the end of force. This delay varied with locomotor speedin the lateral gastrocnemius (LG); it was longer at slow walkingspeeds than for running. This variation in r-EMD was not explainedby differences in muscle length trajectory, as the magnitudeof r-EMD was not correlated with the velocity of shorteningof the muscle during relaxation. We speculate that the longerrelaxation times at slow walking speeds compared with runningmay reflect the longer time course of relaxation in slower musclesfibers. We also examined the relationship between magnitudeof force and EMG across a range of walking and running speeds.We analyzed the force–EMG relationship during the swingphase separately from the force–EMG relationship duringstance phase. During stance, force amplitude (average force)was linearly related to mean EMG amplitude (average EMG). Forcesduring swing phase were lower than predicted from the stancephase force–EMG relationship. The different force–EMGrelationships during the stance and swing phases may reflectthe contribution of passive structures to the development offorce, or a nonlinear force–EMG relationship at low levelsof muscle activity. Together the results suggest that any inferenceof force from EMG must be done cautiously when a broad rangeof activities is considered.     

EMG spectral shift as an indicator of fatigability in an heterogeneous muscle group

Changes in electromyogram (EMG) power spectra were investigated in the triceps surae muscles of two classes of individuals (untrained subjects and athletes) maintaining a plantarflexion torque of 80% of maximal voluntary contraction until exhaustion. A set of 23 parameters describing changes in the frequency content and power of EMG was defined. For most experiments, classical changes were found, indicating a shift of the EMG spectra towards lower frequencies and an increase in the total power of the signals. In 12% of the experiments, alternations in activity between synergistic muscles were found, leading to a large variability in the spectral parameters. After the expression of each experiment in terms of a reduced data matrix and matrix to vector transformations, three methods of discrimination were used to classify subjects with respect to changes in the EMG signal during sustained contraction: (1) evaluation of the most discriminating parameter, (2) principal components analysis, (3) transformation maximizing differences between classes. Method (3) was found to be preferable since it led to good separation of the two classes in a reference group of subjects and a satisfactory projection of each individual from a group of unknowns into the appropriate class. These results suggest using a method such as this for ergonomic or athletic training purposes rather than the usual method of monitoring the frequency shift of the EMG.     

Comparison between EMG to force processing and kinetic analysis for the calf muscle moment in walking and stepping

In EMG to force processing the force of a muscle or the corresponding muscle moment with respect to the joint is determined from EMG and muscle length by means of a model of the muscle's contractile and elastic properties. In kinetic analysis the joint moment is calculated from the ground reaction force and the positions and accelerations of the body segments by means of Newtonian equations, based on a rigid body segment model. These two fundamentally different methods have been compared by determining simultaneously the calf muscle moment and the total ankle moment in walking at slow, moderate and fast speed and in stepping up and down a low bench. As long as there is no activity of other muscles, the moments obtained by either method should be identical. The analysis was restricted to such periods, as assessed from the EMG of tibialis anterior. In each experimental condition three steps were analyzed in five subjects of 21-22 yr. In comparison with the results of kinetic analysis, EMG processing shows differences between 7 and 54 Nm r.m.s., with an average of 22 Nm r.m.s. Normalized with respect to the r.m.s. value of the moment itself, the differences amount to 0.13-0.51, with an average of 0.22. In the majority of cases there was no systematic deviation between EMG processing and kinetic analysis and the r.m.s. difference was of the same order of magnitude as determined previously in ergometer experiments.     

A biomechanical model for analysis of muscle force,power output and lower jaw motion in fishes

Fish skulls are complex kinetic systems with movable components that are powered by muscles. Cranial muscles for jaw closing pull the mandible around a point of rotation at the jaw joint using a third-order lever mechanism. The present study develops a lever model for the jaw of fishes that uses muscle design and the Hill equation for nonlinear length-tension properties of muscle to calculate dynamic power output. The model uses morphometric data on skeletal dimensions and muscle proportions in order to predict behavior and force transmission mediated by lever action. The computer model calculates a range of dynamic parameters of jaw function including muscle force, torque, effective mechanical advantage, jaw velocity, bite duration, bite force, work and power. A complete list of required morphometrics is presented and a software program (MandibLever 2.0) is available for implementing lever analysis. Results show that simulations yield kinematics and timing profiles similar to actual fish feeding events. Simulation of muscle properties shows that mandibles reach their peak velocity near the start of jaw closing, peak force at the end of jaw closing, and peak power output at about 25% of the closing cycle time. Adductor jaw muscles with different mechanical designs must have different contractile properties and/or different muscle activity patterns to coordinate jaw closing. The effective mechanical advantage calculated by the model is considerably lower than the mechanical advantage estimated from morphological lever ratios, suggesting that previous studies of morphological lever ratios have overestimated force and underestimated velocity transmission to the mandible. A biomechanical model of jaw closing can be used to interpret the mechanics of a wide range of jaw mechanisms and will enable studies of the functional results of developmental and evolutionary changes in skull morphology and physiology.     

A preliminary analysis of EMG variance as an index of change in EMG biofeedback treatment of tension-type headache

The effectiveness of EMG biofeedback training for tension headache has been well established. Previous studies evaluating changes in an average EMG activity score from pre- to posttreatment have not consistently found a relationship between a reduction in average EMG activity and headache improvement at posttreatment. The current study is a preliminary analysis of the utility of EMG variance as another possible mechanism of change. Frontalis EMG average activity and variances from 6 chronic tension-type headache sufferers who demonstrated significant improvement in headache activity at posttreatment (at least 70%) and 6 chronic tension-type headache sufferers who did not demonstrate improvement (less than 30%) were examined across 6 sessions of biofeedback treatment. The improved group demonstrated larger time-specific EMG variance in relation to mean EMG amplitudes during all treatment sessions. A dramatic decline in time-specific variance was observed during the later treatment sessions for improved participants; this pattern was not observed in the group who demonstrated little or no improvement. Results from the current study suggest that the inclusion of both average EMG activity and EMG variance may provide a more comprehensive measure to evaluate possible physiological changes responsible for improvement in headache activity following EMG biofeedback training.     

A feasibility study for the use of a silastic gage as an in vivo muscle force transducer

This study investigates the feasibility of utilizing silastic gages for in vivo dynamic muscle force measurement. The gastrocnemius muscle of a fifty-one pound black short hair dog was selected for the test. The study shows that such measurements can be reliably performed in vivo for short durations without interfering with the natural movement of the animal. The durability of the gage appears to be primarily limited by the biological rejection process at the gage site.     

Epidemic pasteurellosis in a bighorn sheep population coinciding with the appearance of a domestic sheep

A pneumonia epidemic reduced bighorn sheep (Ovis canadensis) survival and recruitment during 1997-2000 in a population comprised of three interconnected wintering herds (Kenosha Mountains, Sugarloaf Mountain, Twin Eagles) that inhabited the Kenosha and Tarryall Mountain ranges in central Colorado, USA. The onset of this epidemic coincided temporally and spatially with the appearance of a single domestic sheep (Ovis aires) on the Sugarloaf Mountain herd's winter range in December 1997. Although only bighorns in the Sugarloaf Mountain herd were affected in 1997-98, cases also occurred during 1998-99 in the other two wintering herds, likely after the epidemic spread via established seasonal movements of male bighorns. In all, we located 86 bighorn carcasses during 1997-2000. Three species of Pasteurella were isolated in various combinations from affected lung tissues from 20 bighorn carcasses where tissues were available and suitable for diagnostic evaluation; with one exception, beta-hemolytic mannheimia (Pasteurella) haemolytica (primarily reported as biogroup 1(G) or 1(alphaG)) was isolated from lung tissues of cases evaluated during winter 1997-98. The epidemic dramatically lowered adult bighorn monthly survival in all three herds; a model that included an acute epidemic effect, differing between sexes and with vaccination status, that diminished linearly over the next 12 mo best represented field data. In addition to the direct mortality associated with epidemics in these three herds, lamb recruitment in years following the pneumonia epidemic also was depressed as compared to years prior to the epidemic. Based on observations presented here, pasteurellosis epidemics in free-ranging bighorn sheep can arise through incursion of domestic sheep onto native ranges, and thus minimizing contact between domestic and bighorn sheep appears to be a logical principle for bighorn sheep conservation.     

EMG, muscle fibre and force production characteristics during a 1 year training period in elite weight-lifters

The effects of a 1 year training period on 13 elite weight-lifters were investigated by periodical tests of electromyographic, muscle fibre and force production characteristics. A statistically non-significant increase of 3.5% in maximal isometric strength of the leg extensors, from 4841 +/- 1104 to 5010 +/- 1012 N, occurred over the year. Individual changes in the high force portions of the force-velocity curve correlated (p less than 0.05-0.01) with changes in weight-lifting performance. Training months 5-8 were characterized by the lowest average training intensity (77.1 +/- 2.0%), and this resulted in a significant (p less than 0.05) decrease in maximal neural activation (IEMG) of the muscles, while the last four month period, with only a slightly higher average training intensity (79.1 +/- 3.0%), led to a significant (p less than 0.01) increase in maximum IEMG. Individual increases in training intensity between these two training periods correlated with individual increases both in muscular strength (p less than 0.05) and in the weight lifted in the clean & jerk (p less than 0.05). A non-significant increase of 3.9% in total mean muscle fibre area occurred over the year. The present findings demonstrate the limited potential for strength development in elite strength athletes, and suggest that the magnitudes and time courses of neural and hypertrophic adaptations in the neuromuscular system during their training may differ from those reported for previously untrained subjects. The findings additionally indicate the importance of training intensity for modifying training responses in elite strength athletes.     

The influence of an increase in the level of force on the EMG power spectrum of elbow extensors

It has been proposed that the mean power frequency (MPF) of the electromyogram (EMG) power spectrum increases gradually with force of contraction and that this increase is a function of the fiber-type content of the muscle investigated and the inter-electrode distance (IED) used when recording the EMG signals. In order to test these hypotheses, the values of the MPF of two elbow extensor muscles, triceps brachii (TB, 65% fast twitch fibers) and anconeus (AN, 65% slow twitch fibers), were compared at different levels of contraction. Subjects (n = 13) produced ten static ramp elbow extensions [0-100% maximum voluntary contraction (MVC)]. EMG signals of each muscle were recorded with two pairs of surface miniature electrodes having IEDs of 6 mm and 30 mm respectively. MPFs were obtained at each of the following levels: 10, 20, 40, 60, 80 and 100% MVC. Statistical analyses indicated that the MPF of AN increased significantly (P less than 0.05) up to 60% MVC. In contrast, the MPF values for TB showed no significant change across different levels of contraction (P greater than 0.05). Since skinfold was on average 3.2 times thicker over TB than over AN it is suggested that the low-pass filtering effect of the skin could have prevented the observation of an increase of the MPF for TB. It thus appears that changes of the MPF with the level of force, as disclosed by surface electrode recordings, is specific to each muscle. Consequently one has to account for factors such as thickness of the skinfold when it comes to the determination of the fiber-type content of different muscles within a subject.     

Effects of EMG processing on biomechanical models of muscle joint systems: sensitivity of trunk muscle moments, spinal forces, and stability

Biomechanical models are in use to estimate parameters such as contact forces and stability at various joints. In one class of these models, surface electromyography (EMG) is used to address the problem of mechanical indeterminacy such that individual muscle activation patterns are accounted for. Unfortunately, because of the stochastical properties of EMG signals, EMG based estimates of muscle force suffer from substantial estimation errors. Recent studies have shown that improvements in muscle force estimation can be achieved through adequate EMG processing, specifically whitening and high-pass (HP) filtering of the signals. The aim of this paper is to determine the effect of such processing on outcomes of a biomechanical model of the lumbosacral joint and surrounding musculature. Goodness of fit of estimated muscle moments to net moments and also estimated joint stability significantly increased with increasing cut-off frequencies in HP filtering, whereas no effect on joint contact forces was found. Whitening resulted in moment estimations comparable to those obtained from optimal HP filtering with cut-off frequencies over 250 Hz. Moreover, compared to HP filtering, whitening led to a further increase in estimated joint-stability. Based on theoretical models and on our experimental results, we hypothesize that the processing leads to an increase in pick-up area. This then would explain the improvements from a better balance between deep and superficial motor unit contributions to the signal.