Simulation of Respiratory Mechanics

The dynamic relationship between respiratory muscle effort and the consequent changes in lung volume is investigated. A mathematical simulation based on the structures that form the connection between these two variables makes it possible to lump the contribution from all respiratory muscles into a single time-varying driving force. When this force is applied to the system model, the appropriate lung volume pattern results. The simulation results indicate the accuracy of the model and the validity of the lumped muscle force assumption. In addition, the system model adequately describes abnormal conditions such as decreased lung compliance and increased airway resistance. The results of this simulation suggest that the modeling technique is extremely useful in describing and analyzing complex respiratory system interactions.     

Eccentric and posteccentric contractile behaviour of skeletal muscle a comparative study in frog single fibres and in humans

Eccentric and posteccentric force behaviour in human skeletal muscle and in isolated frog muscle fibres was studied by imposing stretch-and-hold loading conditions during contractions with maximal voluntary effort or under tetanic stimulation in the isolated preparations. The investigations on human muscle were made on the forearm flexors of a group of kayak racers (n = 16; age: 17-22 years) and of schoolgirls (n = 15; age: 17-18 years) with both groups participating in a strength-training programme over 4 (kayak racers) or 3 (girls) months. Half of the training regime consisted of eccentric elements. In the isolated muscle fibres, it could be shown that in the posteccentric hold phase the enhanced force decayed exponentially to the original isometric value with a mean time-constant of 0.35 s (10 degrees C) and of 0.23 (20 degrees C). In the forearm flexor of human subjects similar results were obtained not only qualitatively but even quantitatively (time constant of posteccentric force decay: 0.25-0.37 s). Strength training in both groups did not lead to an enhancement in maximal isometric force alone [mean increase in force 17 (SD 10)%], a well-known and generally accepted fact, but also to a parallel shift in eccentric [21 (SD 10)%] and posteccentric force level. The close similarity between the findings in isolated muscle fibres and in human muscle in situ suggests that the eccentric and posteccentric behaviour must be primarily ascribed to the contractile properties of the muscle fibres themselves. A three-element muscle model with variable visco-elastic properties would appear to be most suitable for simulating the experimental findings.     

Trunk stiffness increases with steady-state effort

Trunk stiffness was measured in healthy human subjects as a function of steady-state preload efforts in different horizontal loading directions. Since muscle stiffness increases with increased muscle activation associated with increasing effort, it is believed that coactivation of muscles helps to stiffen and stabilize the trunk. This paper tested whether increased steady-state preload effort increases trunk stiffness. Fourteen young healthy subjects each stood in an apparatus with the pelvis immobilized. They were loaded horizontally at directions of 0, 45, 90, 135 and 180 degrees to the forward direction via a thoracic harness. Subjects first equilibrated with a steady-state load of 20 or 40% of their maximum extension effort. Then a sine-wave force perturbation of nominal amplitude of 7.5 or 15% of maximum effort and nominal period of 250ms was applied. Both the applied force and subsequent motion were recorded. Effective trunk mass and trunk-driving point stiffness were estimated by fitting the experimental data to a second-order differential equation of the trunk dynamic behavior. The mean effective trunk mass was 14.1kg (s.d.=4.7). The trunk-driving point stiffness increased on average 36.8% (from 14.5 to 19.8N/mm) with an increase in the nominal steady-state preload effort from 20 to 40% (F(1,13)=204.96, p<0.001). There was a smaller, but significant variation in trunk stiffness with loading direction. The measured increase in trunk stiffness probably results from increased muscle stiffness with increased muscle activation at higher steady-state efforts.     

A dynamical model of muscle activation, fatigue, and recovery

A dynamical model is presented as a framework for muscle activation, fatigue, and recovery. By describing the effects of muscle fatigue and recovery in terms of two phenomenological parameters (F, R), we develop a set of dynamical equations to describe the behavior of muscles as a group of motor units activated by voluntary effort. This model provides a macroscopic view for understanding biophysical mechanisms of voluntary drive, fatigue effect, and recovery in stimulating, limiting, and modulating the force output from muscles. The model is investigated under the condition in which brain effort is assumed to be constant. Experimental validation of the model is performed by fitting force data measured from healthy human subjects during a 3-min sustained maximal voluntary handgrip contraction. The experimental results confirm a theoretical inference from the model regarding the possibility of maximal muscle force production, and suggest that only 97% of the true maximal force can be reached under maximal voluntary effort, assuming that all motor units can be recruited voluntarily. The effects of different motor unit types, time-dependent brain effort, sources of artifacts, and other factors that could affect the model are discussed. The applications of the model are also discussed.     

Perceptions of effort and heaviness during fatigue and during the size-weight illusion

Previous work has shown that force perception and the sense of motor effort are different attributes of sensorimotor function. This study explores the hypothesis that one reason force and effort perceptions are distinct is to inform an individual of impaired motor function when muscular force lags effort. This hypothesis predicts that effort and force perceptions will dissociate when motor function is impaired by fatigue but not during the size-weight illusion. All subjects reported a distinct increase in effort when lifting a standard test weight as fatigue developed. When fatigue was sufficiently marked so that they could barely lift the test weight, they rated their effort as similar to that required to lift a maximal weight in the unfatigued state. The perceived heaviness of the test weight also increased as fatigue developed, but this fatigue-weight illusion was smaller than the increase in effort for all subjects and displayed greater variability. In contrast, both the perceived weight of a small object and the effort required to lift it increased in parallel when small and large objects were lifted sequentially. The size-weight and size-effort illusions appear to be examples of a common phenomenon in which perceptual experience is rescaled to maintain acuity under different working conditions. The fatigue-weight illusion also has the effect of increasing perceptual acuity as the subject's weight lifting range decreases due to fatigue.     

Validation of a biodynamic model of pushing and pulling.

Pushing and pulling during manual material handling can increase the compressive forces on the lumbar disc region while creating high shear forces at the shoe-floor interface. A sagittal plane dynamic model derived from previous biomechanical models was developed to predict L5/S1 compressive force and required coefficients of friction during dynamic cart pushing and pulling. Before these predictions could be interpreted, however, it was necessary to validate model predictions against independently measured values of comparable quantities. This experiment used subjects of disparate stature and body mass, while task factors such as cart resistance and walking speed were varied. Predicted ground reaction forces were compared with those measured by a force platform, with correlations up to 0.67. Predicted erector spinae and rectus abdominus muscle forces were compared with muscle forces derived from RMS-EMGs of the respective muscle groups, using a static force build-up regression relationship to transform the dynamic RMS-EMGs to trunk muscle forces. Although correlations were low, this was attributed in part to the use of surface EMG on subjects of widely varied body mass. The biodynamic model holds promise as a tool for analysis of actual industrial pushing and pulling tasks, when carefully applied.     

Estimating the muscle forces generated in the human lower extremity when walking: a physiological solution

A model capable of estimating individual muscle activity during human lower extremity activity has been developed. This model was implemented and applied to normal gait. Both 3D cinematography and force platform records were collected to define the kinematics of the lower limb segments and the ground reaction forces. From these data the lengths, velocities, and moment arms of the muscles and the net muscle moments were calculated. These data were then used to estimate the output from a set of pattern generators which specified the neural input to and hence the force generated in each muscle. Results were obtained which demonstrate that it is feasible to construct a hierarchical physiological model which will estimate individual muscle forces. This statement must be tempered somewhat in that better anatomical and neurophysiological data must be made available and that this model, and all others attempting to estimate individual muscle forces, must be rigorously tested before being applied in a research, sports, or clinical setting.     

A new model for calculating muscle forces from electromyograms

A muscle model is described that uses electromyogram (EMG), muscle length and speed of contraction to predict muscle force. Physiological parameters are the Hill constants and the shape of the twitch response to a single stimulus. The model was incorporated in a jaw model of the rabbit and tested by predicting the bite force produced by the jaw muscles during mastication. The time course of the calculated force appeared to match the bite force, measured in vivo by a strain gauge, applied to the bone below the teeth. The variation in peak strain amplitude from cycle to cycle correlated with the variation predicted by the model. The peak amplitude of the integrated EMGs of individual jaw muscles showed an average correlation with peak strain of 0.41. Use of the sum of the available peak amplitudes, weighted according to their effect upon the bite force increased the correlation to 0.46; the model predicted bite forces showed a correlation of 0.57 with the strain. The increase in correlation was statistically significant. The muscle forces were calculated using a minimum number of easily obtainable constants.     

Distribution of calcium during contraction and relaxation of crayfish skeletal muscle fibre.

A model of activation of muscle contraction has been applied to the crayfish isolated skeletal muscle fibre. The model is based on calcium diffusion and binding to specific regulatory sites in a sarcomere. Calcium ions activate interactions of contractile proteins and thus the generation of force. The model quantifies the relation between calcium released from intracellular stores and force elicited. Experimental tension records from isolated crayfish skeletal muscle fibres under voltage clamp conditions are analyzed. Model parameters were determined either via approximation of the onset of tension by the model solution or from the model based relations between the tension maximum, and depolarizing pulse length and amplitude. This allowed to determine time changes of free and bound calcium distribution in the sarcomere and the calcium release from terminal cisternae. The steady state calcium concentration at terminal cisternae showed S-shaped voltage dependence with saturation below approx. 10 mumol/l at positive membrane potentials.     

The image of motor units architecture in the mechanomyographic signal during the single motor unit contraction: in vivo and simulation study

The mechanomyographic (MMG) signal analysis has been performed during single motor unit (MU) contractions of the rat medial gastrocnemius muscle. The MMG has been recorded as a muscle surface displacement by using a laser distance sensor. The profiles of the MMG signal let to categorize these signals for particular MUs into three classes. Class MMG-P (positive) comprises MUs with the MMG signal similar to the force signal profile, where the distance between the muscle surface and the laser sensor increases with the force increase. The class MMG-N (negative) has also the MMG profile similar to the force profile, however the MMG is inverted in comparison to the force signal and the distance measured by using laser sensor decreases with the force increase. The third class MMG-M (mixed) characterize the MMG which initially increases with the force increases and when the force exceeds some level it starts to decrease towards the negative values. The semi-pennate muscle model has been proposed, enabling estimation of the MMG generated by a single MU depending on its localization. The analysis have shown that in the semi-pennate muscle the localization of the MU and the relative position of the laser distance sensor determine the MMG profile and amplitude. Thus, proposed classification of the MMG recordings is not related to the physiological types of MUs, but only to the MU localization and mentioned sensor position. When the distance sensor is located over the middle of the muscle belly, a part of the muscle fibers have endings near the location of the sensor beam. For the MU MMG of class MMG-N the deflection of the muscle surface proximal to the sensor mainly influences the MMG recording, whereas for the MU MMG class MMG-P, it is mainly the distal muscle surface deformation. For the MU MMG of MMG-M type the effects of deformation within the proximal and distal muscle surfaces overlap. The model has been verified with experimental recordings, and its responses are consistent and adequate in comparison to the experimental data.     

Effect of muscle fatigue on internal model formation and retention during reaching with the arm.

The motor system adapts to novel dynamic environments by forming internal models that predict the muscle forces needed to move skillfully. The goal of this study was to determine how muscle fatigue affects internal model formation during arm movement and whether an internal model acquired while fatigued could be recalled accurately after rest. Twelve subjects adapted to a viscous force field applied by a lightweight robot as they reached to a target. They then reached while being resisted by elastic bands until they could no longer touch the target. This protocol reduced the strength of the muscles used to resist the force field by approximately 20%. The bands were removed, and subjects adapted again to the viscous force field. Their adaptive ability, quantified by the amount and time constant of adaptation, was not significantly impaired following fatigue. The subjects then rested, recovering approximately 70% of their lost force-generation ability. When they reached in the force field again, their prediction of the force field strength was different than in a nonfatigued state. This alteration was consistent with the use of a higher level of effort than normally used to counteract the force field. These results suggest that recovery from fatigue can affect recall of an internal model, even when the fatigue did not substantially affect the motor system's ability to form the model. Recovery from fatigue apparently affects recall because the motor system represents internal models as a mapping between effort and movement and relies on practice to recalibrate this mapping.     

Trunk muscle activation and associated lumbar spine joint shear forces under different levels of external forward force applied to the trunk.

High anterior intervertebral shear loads could cause low back injuries and therefore the neuromuscular system may actively counteract these forces. This study investigated whether, under constant moment loading relative to L3L4, an increased externally applied forward force on the trunk results in a shift in muscle activation towards the use of muscles with more backward directed lines of action, thereby reducing the increase in total joint shear force. Twelve participants isometrically resisted forward forces, applied at several locations on the trunk, while moments were held constant relative to L3L4. Surface EMG and lumbar curvature were measured, and an EMG-driven muscle model was used to calculate compression and shear forces at all lumbar intervertebral joints. Larger externally applied forward forces resulted in a flattening of the lumbar lordosis and a slightly more backward directed muscle force. Furthermore, the overall muscle activation increased. At the T12L1 to L3L4 joint, resulting joint shear forces remained small (less than 200N) because the average muscle force pulled backward relative to those joints. However, at the L5S1 joint the average muscle force pulled the trunk forward so that the increase in muscle force with increasing externally applied forward force caused a further rise in shear force (by 102.1N, SD=104.0N), resulting in a joint shear force of 1080.1N (SD=150.4N) at 50Nm moment loading. It is concluded that the response of the neuromuscular system to shear force challenges tends to increase rather than reduce the shear loading at the lumbar joint that is subjected to the highest shear forces.     

Exercise-induced respiratory muscle fatigue: implications for performance.

It is commonly held that the respiratory system has ample capacity relative to the demand for maximal O(2) and CO(2) transport in healthy humans exercising near sea level. However, this situation may not apply during heavy-intensity, sustained exercise where exercise may encroach on the capacity of the respiratory system. Nerve stimulation techniques have provided objective evidence that the diaphragm and abdominal muscles are susceptible to fatigue with heavy, sustained exercise. The fatigue appears to be due to elevated levels of respiratory muscle work combined with an increased competition for blood flow with limb locomotor muscles. When respiratory muscles are prefatigued using voluntary respiratory maneuvers, time to exhaustion during subsequent exercise is decreased. Partially unloading the respiratory muscles during heavy exercise using low-density gas mixtures or mechanical ventilation can prevent exercise-induced diaphragm fatigue and increase exercise time to exhaustion. Collectively, these findings suggest that respiratory muscle fatigue may be involved in limiting exercise tolerance or that other factors, including alterations in the sensation of dyspnea or mechanical load, may be important. The major consequence of respiratory muscle fatigue is an increased sympathetic vasoconstrictor outflow to working skeletal muscle through a respiratory muscle metaboreflex, thereby reducing limb blood flow and increasing the severity of exercise-induced locomotor muscle fatigue. An increase in limb locomotor muscle fatigue may play a pivotal role in determining exercise tolerance through a direct effect on muscle force output and a feedback effect on effort perception, causing reduced motor output to the working limb muscles.     

A 3D biomechanical model of the hand for power grip

A three-dimensional scalable biomechanical model of the four fingers of the hand to evaluate power grip is proposed. The model has been validated by means of reproducing an experiment in which the subjects exerted the maximal voluntary grasping force over cylinders of different diameters. The model is used to simulate the cylinder grip for two hand sizes and for five different handle diameters. The reduction of the muscle forces using different handle diameters has been studied. The model can be applied to the design and evaluation of handles for power grip and to the study of power grasp for normal and abnormal hands.     

Effect of abdominal compression on maximum transdiaphragmatic pressure

Transdiaphragmatic pressure (Pdi) is lower during maximum inspiratory effort with the diaphragm alone than when maximum inspiratory and expulsive efforts are combined. The increase in Pdi with expulsive effort has been attributed to increased neural activation of the diaphragm. Alternatively, the increase could be due to stretching of the contracted diaphragm. If this were so, Pdi measured during a combined maximum effort would overestimate the capacity of the diaphragm to generate inspiratory force. This study determined the likely contribution of stretching of the contracted diaphragm to estimates of maximum Pdi (Pdimax) obtained during combined inspiratory and expulsive effort. Three healthy trained subjects were studied standing. Diaphragmatic Mueller maneuvers were performed at functional residual capacity and sustained during subsequent abdominal compression by either abdominal muscle expulsive effort or externally applied pressure. Measurements were made of changes in abdominal (Pab) and pleural (Ppl) pressure, Pdi, rib cage and abdominal dimensions and respiratory electromyograms. Three reproducible performances of each maneuver from each subject were analyzed. When expulsive effort was added to maximum diaphragmatic inspiratory effort, Pdimax increased from 86 +/- 12 to 148 +/- 14 (SD) cmH2O within the 1st s and was 128 +/- 14 cmH2O 2 s later. When external compression was added to maximum diaphragmatic inspiratory effort, Pdimax increased from 87 +/- 16 to 171 +/- 19 cmH2O within the 1st s and was 152 +/- 16 cmH2O 2 s later.(ABSTRACT TRUNCATED AT 250 WORDS)     

Vigilance Decreases with Time at Loafing Sites in Gulls (Larus spp.)

Understanding how animals partition effort between vigilance for predators and other conflicting activities has been a mainstay of animal behaviour research. Classical theories implicitly assume that animals alternate between high and low vigilance states over short timescales, but that average effort invested in vigilance is constant over an extended bout of such alternations. However, one recent model suggests that vigilance should be adjusted dynamically to short‐term changes in the perception of predation risk and would tend to decrease with time. Indeed, as time passes by without disturbances, perception of the need for vigilance should decrease and prey animals may allocate more time to competing activities. Here, we examined how the proportion of sleeping gulls (Larus spp.) in loafing groups changed over time. Sleeping gulls can only maintain low levels of vigilance against external threats (compared to alert individuals), and we predicted that the proportion of sleeping gulls at loafing sites should increase over time when no disturbances occur. Statistically significant changes in the proportion of sleeping gulls as a function of time occurred in the majority of sequences and an increase was observed significantly more often than predicted by chance alone. This temporal pattern cannot be caused by reduction in hunger levels because gulls are not feeding at loafing sites. The results indicate that vigilance can be adjusted dynamically in response to short‐term temporal changes in the perception of predation risk.     

Passive force characteristics of an architecturally complex muscle

In architecturally complex muscles with large attachment areas, it can be expected that during movement different muscle regions undergo different amounts of length excursions. As a consequence, the amount of passive force produced by the regions will differ. Therefore, we tested the hypothesis that during movement the vector of the passive force of such a muscle, which defines the magnitude, position and orientation of the resultant force of the various regions, has no fixed position, between the muscle's center of origin and insertion. As a model for an architecturally complex muscle we used the masseter muscle. It was expected that during jaw opening anterior muscle regions are more stretched than posterior regions, leading to an anterior shift of the passive force vector. A three-component force transducer was used to measure both the position and magnitude of passive force in the masseter muscle of 9 rabbits. Forces were recorded during repeated cycles of stepwise opening and closure of the jaw. The muscle exhibited a clear hysteresis: passive force measured during jaw opening was larger than that during jaw closing. With an increase of the jaw gape there was an approximately exponential increase of the magnitude of the passive muscle force, while simultaneously the passive force vector shifted anteriorly. Moment arm length of passive force increased by about 100%. This anterior shift contributed substantially to the increase of the passive muscle moment generated during jaw opening. It can be concluded that in architecturally complex muscles the increase of the passive resistance moment which is associated with muscle lengthening might not only be due to an increase of the magnitude of passive muscle force but also to an increase of the moment arm of this force.     

Cross-bridge mechanism of residual force enhancement after stretching in a skeletal muscle

A muscle model that uses a modified Langevin equation with actomyosin potentials was used to describe the residual force enhancement after active stretching. Considering that the new model uses cross-bridge theory to describe the residual force enhancement, it is different from other models that use passive stretching elements. Residual force enhancement was simulated using a half sarcomere comprising 100 myosin molecules. In this paper, impulse is defined as the integral of an excess force from the steady isometric force over the time interval for which a stretch is applied. The impulse was calculated from the force response due to fast and slow muscle stretches to demonstrate the viscoelastic property of the cross-bridges. A cross-bridge mechanism was proposed as a way to describe the residual force enhancement on the basis of the impulse results with reference to the compliance of the actin filament. It was assumed that the period of the actin potential increased by 0.5% and the amplitude of the potential decreased by 0.5% when the half sarcomere was stretched by 10%. The residual force enhancement after 21.0% sarcomere stretching was 6.9% of the maximum isometric force of the muscle; this value was due to the increase in the number of cross-bridges.     

The effect of hamstring muscle compensation for anterior laxity in the ACL-deficient knee during gait

The hamstring muscles have been recognized as an important element in compensating for the loss of stability in the ACL-deficient knee, but it is still not clear whether the hamstring muscle force can completely compensate for the loss of ACL, and the consequences of increased hamstring muscle force. A two-dimensional anatomical knee model in the sagittal plane was developed to examine the effect of various levels of hamstring muscle activation on restraining anterior tibial translation in the ACL-deficient knee during level walking. The model included the tibiofemoral and patellofemoral joints, four major ligaments, the medial capsule, and five muscle units surrounding the knee. Simulations were conducted to determine anterior tibial translation and internal joint loading at a single selected position when the knee was under a peak external flexion moment during early stance phase of gait. Incremental hamstring muscle forces were applied to the modeled normal and the ACL-deficient knees. Results of simulations showed that the ACL injury increased the anterior tibial translation by 11.8mm, while 56% of the maximal hamstring muscle force could reduce the anterior translation of the tibia to a normal level during the stance phase of gait. The consequences of increased hamstring muscle force included increased quadriceps muscle force and joint contact force.     

Extrapolation of an empirical elbow muscle co-activation relationship to a novel task set: implications for predictions of individual muscle demands

Biomechanical optimisation models applying efficiency-based objective functions often underestimate antagonist contributions. Previous work has quantified an empirical co-activation relationship in the elbow musculature, demonstrating that implementing this relationship as a constraint in an elbow muscle force prediction model improves muscle force predictions. The current study evaluated this modified model by extrapolating the co-activation relationship to 36 novel isometric unilateral, right-handed exertions, including those requiring greater intensity of effort and performed in different postures. Surface electromyography was recorded from the elbow flexors and extensors. Novel extrapolative co-activation relationships were developed and used as constraints in a muscle force prediction model. Model predictions using both constraints were compared with empirical biophysical data. Predictions by the modified model were more consistent with biophysical data than those by the original model for the novel exertions. Novel co-activation relationships did not further enhance predictions when compared with the previous relationship, suggesting that extrapolation of the previous relationship is feasible.