Trunk response to sudden forward perturbations – Effects of preload and sudden load magnitudes,posture and abdominal antagonistic activation

Unexpected loading of the spine is a risk factor for low back pain. The trunk neuromuscular and kinematics responses are likely influenced by the perturbation itself as well as initial trunk conditions. The effect of four parameters (preload, sudden load, initial trunk flexed posture, initial abdominal antagonistic activity) on trunk kinematics and back muscles reflex response were evaluated. Twelve asymptomatic subjects participated in sudden forward perturbation tests under six distinct conditions. Preload did not change the reflexive response of back muscles and the trunk displacement; while peak trunk velocity and acceleration as well as the relative load peak decreased. Sudden load increased reflex response of muscles, trunk kinematics and loading variables. When the trunk was initially flexed, back muscles latency was delayed, trunk velocity and acceleration increased; however, reflex amplitude and relative trunk displacement remained unchanged. Abdominal antagonistic preactivation increased reflexive response of muscles but kinematics variables were not affected. Preload, initial flexed posture and abdominal muscles preactivation increased back muscles preactivity. Both velocity and acceleration peaks of the trunk movement decreased with preload despite greater total load. In contrast, they increased in the initial flexed posture and to some extent when abdominal muscles were preactivated demonstrating the distinct effects of pre-perturbation variables on trunk kinematics and risk of injury.     

Muscle dynamics: Dependence of muscle length on changes in external load

A phenomenological theory of muscle dynamics has been elaborated on the basis of data obtained in experiments on hind limb extensor muscles of narcotized cat. Functional dependence of muscle length on external load was explored in conditions of a constant frequency of the efferent stimulation. It was shown that the system under study could be presented for a rather wide class of input signals as a system with nonlinear statics and linear dynamics. The nonlinear statics was shown to be determined mainly by the hysteretical effects of muscle contraction, whereas dynamic element was described by the first order linear differential equation corresponding to the traditional three-component mechanical model of the muscle. A hypothesis was proposed to explain the hysteresis in active muscle on the basis of functioning of the troponin-tropomyosin regulatory complex. Elaborated mathematical model of muscle dynamics can be used to predict and evaluate changes in the muscle length evoked by arbitrary changes in the external load.     

A three-dimensional muscle model: a quantified relation between form and function of skeletal muscles

A three-dimensional muscle model with complex geometry is described and tested against experimental data. Using this model, several muscles were constructed. These muscles have equal optimum length but differ in architecture. The force exerted by the constructed muscles, in relation to their actual length and velocity of shortening, is discussed. Generally speaking, the constructed muscles with considerable pennation have great fiber angles, a great physiological cross section, a narrow active and steep passive length-force relation, and a low maximal velocity of shortening. The maximal power (force times velocity) delivered by the constructed muscles is shown to be almost independent of the architecture of the muscles. The steepness of the passive length-force relation is determined mainly by the shortest fibers within the group of constructed muscles, whereas maximal velocity of shortening and the width of the active length-force relation are determined mainly by the longest fibers. The validity of the three-dimensional muscle model with respect to some morphological and functional characteristics is tested. Length-force relations of constructed muscles are compared with the actual length-force relations of mm. gastrocnemii mediales and mm. semimembranosi of male Wistar rats. Moreover, actual fiber angle, fiber length, and muscle thickness of three mm. gastrocnemii mediales are compared with values found for constructed muscles. It is concluded that the three-dimensional muscle model closely approximates the actual muscle form and function.     

The effect of shortening history on isometric and dynamic muscle function

Despite numerous reports on isometric force depression, few reports have quantified force depression during active muscle shortening (dynamic force depression). The purpose of this investigation was to determine the influence of shortening history on isometric force following active shortening, force during isokinetic shortening, and velocity during isotonic shortening. The soleus muscles of four cats were subjected to a series of isokinetic contractions at three shortening velocities and isotonic contractions under three loads. Muscle excursions initiated from three different muscle lengths but terminated at a constant length. Isometric force produced subsequent to active shortening, and force or shortening velocity produced at a specific muscle length during shortening, were compared across all three conditions. Results indicated that shortening history altered isometric force by up to 5%, force during isokinetic shortening up to 30% and shortening velocity during isotonic contractions by up to 63%. Furthermore, there was a load by excursion interaction during isotonic contractions such that excursion had the most influence on shortening velocity when the loads were the greatest. There was not a velocity by excursion interaction during isokinetic contractions. Isokinetic and isotonic power–velocity relationships displayed a downward shift in power as excursions increased. Thus, to discuss force depression based on differences in isometric force subsequent to active shortening may underestimate its importance during dynamic contractions. The presence of dynamic force depression should be realized in sport performance, motor control modeling and when controlling paralyzed limbs through artificial stimulation.     

A self-organizing model of walking patterns of insects

Insects generate walking patterns which depend upon external conditions. For example, when an insect is exposed to an additional load parallel to the direction in which it is walking, the walking pattern changes according to the magnitude of the load. Furthermore, even after some of its legs have been amputated, an insect will produce walking patterns with its remaining legs. These adaptations in insect walking could not previously be explained by a mathematical model, since the mathemati cal models were based upon the hypothesis that the relationship between walking velocity and walking patterns is fixed under all conditions. We have produced a mathematical model which describes self-organizing insect walking patterns in real-time by using feedback information regarding muscle load (Kimura et al. 1993). As part of this model, we introduced a new rule to coordinate leg movement, in which the information is circulated to optimize the efficiency of the energy transduction of each effector orga n. We describe this mechanism as ‘the least dissatisfaction for the greatest number of elements’. In this paper, we introduce the following aspects of this model, which reflect adaptability to changing circumstances: (1) after one leg is exposed to a transient perturbation, the walking pattern recovers swiftly; (2) when the external load parallel to the walking direction is continuously increased or decreased, the pattern transition point is shifted according to the magnitude of the load increme nt or decrement. This model generates a walking pattern which optimizes energy consumption at a given walking velocity even under these conditions; and (3) when some of the legs are amputated, the model generates walking patterns which are consistent with experimental results. We also discuss the ability of a hierarchical self-organizing model to describe a swift and flexible information processing system. Received: 8 February 1993/Accepted in revised form: 12 November 1993     

Mechanics and energetics of myosin molecular motors from nonpregnant human myometrium

Mechanical properties of spontaneously contracting isolated nonpregnant human myometrium (NPHM) were investigated throughout the whole continuum of load from zero load up to isometry. This made it possible to assess the three-dimensional tension-velocity-length (T-V-L) relationship characterizing the level of contractility and to determine crossbridge (CB) kinetics of myosin molecular motors. Seventy-seven muscle strips were obtained from hysterectomy in 42 nonpregnant patients. Contraction and relaxation parameters were measured during spontaneous mechanical activity. The isotonic tension-peak velocity (T-V) relationship was hyperbolic in 30 cases and nonhyperbolic in 47 cases. When the T-V relationship was hyperbolic, the Huxley formalism could be used to calculate CB kinetics and CB unitary force. At the whole muscle level and for a given isotonic load level, part of the V-L phase plane showed a common pathway, so that a given instantaneous length corresponded to only one possible instantaneous velocity, independent of time and initial length. At the molecular level, rate constants for CB attachment and detachment were dramatically low, ～100 times lower than those of striated muscles, and ～5 to 10 times lower than those of other smooth muscles. The CB unitary force was ～1.4 ± 0.1 pN. NPHM shared similar basic contractile properties with striated muscles, reflected in the three-dimensional T-V-L relationship characterizing the contractile level. Low CB attachment and detachment rate constants made it possible to generate normal CB unitary force and normal muscle tension in NPHM, even though it contracted extremely slowly compared with other muscles.     

Myosin light chain phosphorylation and contractile performance of human skeletal muscle

Twitch tension and maximal unloaded velocity of human knee extensor muscles were studied under conditions of low phosphate content of the phosphorylatable light chains (P-light chains) of myosin and elevated phosphate content, following a 10-s maximal voluntary isometric contraction (MVC). After the MVC, twitch tension was significantly potentiated, with greater potentiation observed at a shorter muscle length (p less than 0.05). The MVC was associated with at least a twofold increase in phosphate content of the fast (LC2F) and two slow (LC2S and LC2S') P-light chains, but this increase was unrelated to muscle length. No significant differences in knee extension velocity were observed between conditions where P-light chains had low or elevated phosphate content. Positive but nonsignificant correlations were noted between the extent of twitch potentiation and phosphate content of individual P-light chains as well as the percentage of type II muscle fibres in vastus lateralis muscle. No significant relationships were determined for myosin light chain kinase activity and either P-light chain phosphorylation or type II fibre percentage. These data suggest that, unlike other mammalian fast muscles, P-light chain phosphorylation of mixed human muscles is not strongly associated with altered contractile performance.     

Role of trunk muscles in generating follower load in the lumbar spine of neutral standing posture

Recently, experimental results have demonstrated that the load carrying capacity of the human spine substantially increases under the follower load condition. Thus, it is essential to prove that a follower load can be generated in vivo by activating the appropriate muscles in order to demonstrate the possibility that the stability of the spinal column could be maintained through a follower load mechanism. The aim of this study was to analyze the coordination of the trunk muscles in order to understand the role of the muscles in generating the follower load. A three-dimensional finite element model of the lumbar spine was developed from T12 to S1 and 117 pairs of trunk muscles (58 pairs of superficial muscles and 59 pairs of deep muscles) were considered. The follower load concept was mathematically represented as an optimization problem. The muscle forces required to generate the follower load were predicted by solving the optimization problem. The corresponding displacements and rotations at all nodes were estimated along with the follower forces, shear forces, and joint moments acting on those nodes. In addition, the muscle forces and the corresponding responses were investigated when the activations of the deep muscles or the superficial muscles were restricted to 75% of the maximum activation, respectively. Significantly larger numbers of deep muscles were involved in the generation of the follower load than the number of superficial muscles, regardless of the restriction on muscle activation. The shear force and the resultant joint moment are more influenced by the change in muscle activation in the superficial muscles. A larger number of deep trunk muscles were activated in order to maintain the spinal posture in the lumbar spine. In addition, the deep muscles have a larger capability to reduce the shear force and the resultant joint moment with respect to the perturbation of the external load or muscle fatigue compared to the superficial muscles.     

Effect of theophylline on the velocity of diaphragmatic muscle shortening

The present study examined the effect of theophylline on the shortening velocity of submaximally activated diaphragmatic muscle (i.e., muscles were activated by the use of a level of stimulation, 50 Hz, within the range of phrenic neural firing frequencies achieved during breathing, whereas maximum activation is achieved at 300 Hz). Experiments were performed in vitro on strips of diaphragmatic muscle obtained from 21 Syrian hamsters. Muscle shortening velocity was assessed during isotonic contractions against a range of afterloads, and Hill's characteristic equation was used to calculate velocity at zero load. In addition, unloaded shortening velocity was also measured by the slack test, i.e., from the time required for muscles to take up slack after a sudden reduction in muscle length. Theophylline (160 mg/l) increased the velocity of muscle shortening against a wide range of external loads (0-14 N/cm2) and increased the extrapolated unloaded velocity of shortening from 6.4 +/- 0.9 to 7.9 +/- 1.1 (SE) lengths/s (P less than 0.01). Theophylline reduced the time required to take up slack for any given step change in muscle length, increasing the unloaded velocity of shortening assessed by the slack test from 7.6 +/- 0.9 to 9.3 +/- 1.1 lengths/s (P less than 0.002). The effect of theophylline on diaphragmatic shortening velocity was evident at concentrations as low as 40 mg/l and increased progressively as theophylline concentrations were increased to 320 mg/l. Theophylline increased the shortening velocity of fatigued as well as fresh muscles.(ABSTRACT TRUNCATED AT 250 WORDS)     

Does the velocity sensitivity of muscle spindles stabilize the stretch reflex?

A model of the stretch reflex based on detailed experimental studies of muscles, muscle receptors and reflex pathways has been analysed. Muscle receptors respond to the velocity and the acceleration of movement, as well as to muscle length. The effect of the velocity sensitivity on the stability of the stretch reflex is considered. It is concluded that the velocity sensitivity can compensate to some extent for the sluggishness of muscles and the delays introduced by reflex pathways. However, the value of velocity sensitivity found experimentally is greater than that required to stabilize the stretch reflex optimally. The velocity sensitivity will have some tendency to produce oscillation, and it is suggested that a small degree of oscillation may be beneficial to linearize muscle properties.     

On the Lacker-Peskin model for muscular contraction

We study Lacker and Peskin's version of A.F. Huxley's model of muscular contraction. The model, which presumes a continuous (and independent) distribution of sites of attachment for a myosin cross-bridge, is presumed to apply to uniformly excited skeletal muscles under conditions of maximal overlap of fibres. We show that given velocity as a function of time there exists a unique force as a function of time, and that the velocity-force relation is continuous and one-to-one. We also suggest minor modifications of the model.     

Trunk response analysis under sudden forward perturbations using a kinematics-driven model

Accurate quantification of the trunk transient response to sudden loading is crucial in prevention, evaluation, rehabilitation and training programs. An iterative dynamic kinematics-driven approach was used to evaluate the temporal variation of trunk muscle forces, internal loads and stability under sudden application of an anterior horizontal load. The input kinematics is hypothesized to embed basic dynamic characteristics of the system that can be decoded by our kinematics-driven approach. The model employs temporal variation of applied load, trunk forward displacement and surface EMG of select muscles measured on two healthy and one chronic low-back pain subjects to a sudden load. A finite element model accounting for measured kinematics, nonlinear passive properties of spine, detailed trunk musculature with wrapping of global extensor muscles, gravity load and trunk biodynamic characteristics is used to estimate the response under measured sudden load. Results demonstrate a delay of ～200 ms in extensor muscle activation in response to sudden loading. Net moment and spinal loads substantially increase as muscles are recruited to control the trunk under sudden load. As a result and due also to the trunk flexion, system stability significantly improves. The reliability of the kinematics-driven approach in estimating the trunk response while decoding measured kinematics is demonstrated. Estimated large spinal loads highlight the risk of injury that likely further increases under larger perturbations, muscle fatigue and longer delays in activation.     

The effects of increasing mandibular load on electrical activity in the mandibular closer muscles during feeding in the desert locust, Schistocerca gregaria

ABSTRACT. Electrical activity from individual mandibular closer muscle fibres of the desert locust Schistocerca gregaria was monitored during three feeding conditions: chewing soft grass, chewing soft grass against an artificial load, and chewing an incompressible grass. Spiking frequency during bursts of activity was found to increase under conditions of increased artificial, and natural load. These increases in spiking frequency occurred over a frequency range much lower than that necessary to produce a maximal tetanic contraction, and therefore correspond to increases in power output by the closer muscles in response to increases in load. A possible mechanism for this is described.     

Muscle activation during maximal voluntary eccentric and concentric knee extension

The aim of this investigation was to study the relationships among movement velocity, torque output and electromyographic (EMG) activity of the knee extensor muscles under eccentric and concentric loading. Fourteen male subjects performed maximal voluntary eccentric and concentric constant-velocity knee extensions at 45, 90, 180 and 360 degrees.s-1. Myoelectric signals were recorded, using surface electrodes, from the vastus medialis, vastus lateralis and rectus femoris muscles. For comparison, torque and full-wave rectified EMG signals were amplitude-averaged through the central half (30 degrees-70 degrees) of the range of motion. For each test velocity, eccentric torque was greater than concentric torque (range of mean differences: 20%-146%, P less than 0.05). In contrast, EMG activity for all muscles was lower under eccentric loading than velocity-matched concentric loading (7%-31%, P less than 0.05). Neither torque output nor EMG activity for the three muscles changed across eccentric test velocities (P greater than 0.05). While concentric torque increased with decreasing velocity, EMG activity for all muscles decreased with decreasing velocity (P less than 0.05). These data suggest that under certain high-tension loading conditions (especially during eccentric muscle actions), the neural drive to the agonist muscles was reduced, despite maximal voluntary effort. This may protect the musculoskeletal system from an injury that could result if the muscle was to become fully activated under these conditions.     

Z band dynamics as a function of sarcomere length and the contractile state of muscle

The Z band in skeletal muscle has two distinct structural states--a relaxed (small square or ss) form and a maximally activated (basket weave or bw) form. We have examined by electron microscopy and optical diffraction Z lattice forms and dimensions and A band spacings in relaxed, tetanized, stretched, and stretched-and-tetanized rat soleus muscle. We have tested the independent contributions of passive load, active tension, and sarcomere length to Z band state. As the A band spacing decreased with increasing load and increasing sarcomere length in the untetanized muscles, the Z lattice remained in the ss form and the Z spacing changed only slightly. Computer-enhanced images from digitized electron micrographs showed that the ss Z lattice resisted deformation regardless of load or method of stretching. In contrast, when the muscle was tetanized at sarcomere lengths of up to 2.7 microns, the Z lattice assumed the bw form and the Z spacing was increased by 20%. Regardless of lattice form, Z spacing did not vary significantly with sarcomere length. Images from freeze-substituted preparations showed both lattice forms comparable to those in images from glutaraldehyde-fixed muscles. Thus, Z band state appears to be a function of the presence (or absence) of active tension. Our previous three-dimensional model is compatible with these observations and with the sub-structures revealed by computer-enhanced images of both lattice forms.     

A kinematic theory of rapid human movements

 This paper proposes a kinematic theory that can be used to study and analyze rapid human movements. It describes a synergy in terms of the agonist and antagonist neuromuscular systems involved in the production of these movements. It is shown that these systems have a log-normal impulse response that results from the limiting behavior of a large number of interdependent neuromuscular networks, as predicted by the central limit theorem. The delta log-normal law that follows from this model is very general and can reproduce almost perfectly the complete velocity patterns of an end-effector. The theory accounts for the invariance and rescalability of these patterns, as well as for the various observations that have been reported concerning the change in maximum and mean velocities, time to maximum velocity, etc., under different experimental conditions. Movement time, load effects, and control strategies are discussed in a companion paper. Received: 15 February 1993/Accepted in revised form: 15 September 1994     

Activation in a Skeletal Muscle Contraction Model with a Modification for Insect Fibrillar Muscle

A sliding filament model for muscle contraction is extended by including an activation mechanism based on the hypothesis that the binding of calcium by a regulating protein in the myofibrils must occur before the rate constant governing the making of interactions between cross-bridges and thin filament sites can take on nonzero values. The magnitude of the rate constant is proportional to the amount of bound calcium. The model's isometric twitch and rise of force in an isometric tetanus are similar to the curves produced by real muscles. It redevelops force after a quick release in an isometric tetanus faster than the initial rise. Quick release experiments on the model during an isometric twitch show that the “active state” curve produced is different from the postulated calcium binding curve. The force developed by the model can be increased by a small quick stretch delivered soon after activation to values near the maximum generated in an isometric tetanus. Following the quick stretch, the force remains near the tetanic maximum for a long time even though the calcium binding curve rises to a peak and subsequently decays by about 50%. The model satisfies the constraint of shortening with a constant velocity under a constant load. Modifications can be made in the model so that it produces the delayed force changes following step length changes characteristic of insect fibrillar muscle.     

Computer simulation of movement-generating cross-bridges.

A stochastic computational method was developed to study properties of cross-bridge models for muscle contraction, by following the time history of individual cross-bridge model of Andrew Huxley (1957) and a modified two-state model with more realistic behavior during steady stretching are used as examples. The method can readily compute steady-state force during shortening and stretching and force-transients following rapid changes in length. Computations of velocity with a steady load and of velocity transients are more sensitive to the randomness inherent in the stochastic method.     

A myoelectrically based dynamic three-dimensional model to predict loads on lumbar spine tissues during lateral bending.

This work describes a dynamic model of the low back that incorporates extensive anatomical detail of the musculo-ligamentous-skeletal system to predict the load time histories of individual tissues. The dynamic reaction moment about L4/L5 was determined during lateral bending from a linked-segment model. This reaction moment was partitioned into restorative components provided by the disc, ligament strain, and active-muscle contraction using a second model of the spine that incorporated a detailed representation of the anatomy. Muscle contraction forces were estimated using both information from surface electromyograms, collected from 12 sites, and consideration of the modulating effects of muscle length, cross-sectional area and passive elasticity. This modelling technique is sensitive to the different ways in which individuals recruit their musculature to satisfy moment constraints. Time histories of muscle forces are provided. High muscle loads are consistent with the common clinical observation of muscle strain often produced by load handling. Furthermore, the coactivation measured in muscles on both sides of the trunk suggests that muscles are recruited to satisfy the lateral bending reaction torque in addition to performing other mechanical roles such as spine stabilization. If an estimate of the intervertebral joint compression is desired for assessment of lateral bends in industry, then a single equivalent lateral muscle with a moment arm of approximately 3.0-4.0 cm would conservatively capture the effects of muscle co-contraction quantified in this study.