Torso flexion modulates stiffness and reflex response.

Neuromuscular factors that contribute to spinal stability include trunk stiffness from passive and active tissues as well as active feedback from reflex response in the paraspinal muscles. Trunk flexion postures are a recognized risk factor for occupational low-back pain and may influence these stabilizing control factors. Sixteen healthy adult subjects participated in an experiment to record trunk stiffness and paraspinal muscle reflex gain during voluntary isometric trunk extension exertions. The protocol was designed to achieve trunk flexion without concomitant influences of external gravitational moment, i.e., decouple the effects of trunk flexion posture from trunk moment. Systems identification analyses identified reflex gain by quantifying the relation between applied force disturbances and time-dependent EMG response in the lumbar paraspinal muscles. Trunk stiffness was characterized from a second order model describing the dynamic relation between the force disturbances versus the kinematic response of the torso. Trunk stiffness increased significantly with flexion angle and exertion level. This was attributed to passive tissue contributions to stiffness. Reflex gain declined significantly with trunk flexion angle but increased with exertion level. These trends were attributed to correlated changes in baseline EMG recruitment in the lumbar paraspinal muscles. Female subjects demonstrated greater reflex gain than males and the decline in reflex gain with flexion angle was greater in females than in males. Results reveal that torso flexion influences neuromuscular factors that control spinal stability and suggest that posture may contribute to the risk of instability injury.     

A new method for measuring passive length-tension properties of human gastrocnemius muscle in vivo

The study of muscle growth and muscle length adaptations requires measurement of passive length-tension properties of individual muscles, but until now such measurements have only been made in animal muscles. We describe a new method for measuring passive length-tension properties of human gastrocnemius muscles in vivo. Passive ankle torque and ankle angle data were obtained as the ankle was rotated through its full range with the knee in a range of positions. To extract gastrocnemius passive length-tension curves from passive torque-angle data it was assumed that passive ankle torque was the sum of torque due to structures which crossed only the ankle joint (this torque was a 6-parameter function of ankle joint angle) and a torque due to the gastrocnemius muscle (a 3-parameter function of knee and ankle angle). Parameter values were estimated with non-linear regression and used to reconstruct passive length-tension curves of the gastrocnemius. The reliability of the method was examined in 11 subjects by comparing three sets of measurements: two on the same day and the other at least a week later. Length-tension curves were reproducible: the average root mean square error was 5.1+/-1.1 N for pairs of measurements taken within a day and 7.3+/-1.2 N for pairs of measurements taken at least a week apart (about 3% and 6% of maximal passive tension, respectively). Length-tension curves were sensitive to mis-specification of moment arms, but changes in length-tension curves were not. The new method enables reliable measurement of passive length-tension properties of human gastrocnemius in vivo, and is likely to be useful for investigation of changes in length-tension curves over time.     

Modelling the line of action for the oblique abdominal muscles using an elliptical torso model

When modelling the line of action of a muscle, anatomical considerations must be included if the model is to realistically mimic the muscle behaviour. The internal and external oblique muscles are examples of muscles that do not follow a straight line between origin and insertion, instead having to wrap around the torso. A model is presented which describes the shape of the torso using a right elliptical cylinder of varying dimensions. The muscle lines of action are then calculated based on this underlying torso shape. The model has been successfully fitted to the data reported by Stokes and Gardner-Morse (Journal of Biomechanics 32(3) (1999) 311). When compared to a linear model, the use of the torso model results in a 15% increase in the axial twist moment, and decreases in the lateral bend and extension moments (5% and 2%, respectively), able to be generated by the internal and external oblique muscles combined in upright stance. These differences become larger (up to 37%) when the torso is flexed, extended or twisted. The structure of the torso model allows it to be used to model any posture without significant increases in the overall model complexity.     

Whole muscle length-tension relationships are accurately modeled as scaled sarcomeres in rabbit hindlimb muscles

An a priori model of the whole active muscle length-tension relationship was constructed utilizing only myofilament length and serial sarcomere number for rabbit tibialis anterior (TA), extensor digitorum longus (EDL), and extensor digitorum II (EDII) muscles. Passive tension was modeled with a two-element Hill-type model. Experimental length-tension relations were then measured for each of these muscles and compared to predictions. The model was able to accurately capture the active-tension characteristics of experimentally-measured data for all muscles (ICC=0.88 ± 0.03). Despite their varied architecture, no differences in predicted versus experimental correlations were observed among muscles. In addition, the model demonstrated that excursion, quantified by full-width-at-half-maximum (FWHM) of the active length-tension relationship, scaled linearly (slope=0.68) with normalized muscle fiber length. Experimental and theoretical FWHM values agreed well with an intraclass correlation coefficient of 0.99 (p<0.001). In contrast to active tension, the passive tension model deviated from experimentally-measured values and thus, was not an accurate predictor of passive tension (ICC=0.70 ± 0.07). These data demonstrate that modeling muscle as a scaled sarcomere provides accurate active functional but not passive functional predictions for rabbit TA, EDL, and EDII muscles and call into question the need for more complex modeling assumptions often proposed.     

Sensory representation of a non‐planar joint in the walking legs of the horseshoe crab,Limulus polyphemus

The patello‐tibial joint in the walking leg of the horseshoe crab can move in the flexion‐extension and promotion‐remotion planes with the freedom of movement of a ball‐and‐socket joint. Its mechanical status is represented by at least seven populations of sensory neurons and these include three different types of proprioceptors; articular membrane receptors, muscle length receptors and muscle tension receptors. A population of each of these types of proprioceptor is associated with structures on the anterior and posterior sides of each leg. In this report we describe the representation of joint movement in both planes by two populations of each type of proprioceptor. In general, the articular membrane receptors were most sensitive to joint flexion; receptors on the anterior side of the leg were biased toward promotion, while those on the posterior were biased toward remotion. Muscle length receptor activity was increased by joint extension and was slightly augmented, in posterior and anterior length receptors, by promotion and remotion, respectively. Length receptor tonic activity was a linear function of muscle length for extension but a non‐linear function of muscle flexion. Tension receptor activity was always highly dependent upon active muscular contraction. Tension receptors associated with posterior tibial flexor showed enhanced activity when the joint was promoted; the activity of those associated with the anterior flexor was enhanced by remotion. The relationships between tension receptor activity and muscle length, and between tension receptor activity and joint position, were also explored. While no single receptor population appears capable of representing the status of this joint in both planes, the output of all of the receptor populations studied appears to provide the CNS with unambiguous information about several parameters representing the status of this complex joint.     

Interaction of viscoelastic tissue compliance with lumbar muscles during passive cyclic flexion–extension

Human and animal models using electromyography (EMG) based methods have hypothesized that viscoelastic tissue properties becomes compromised by prolonged repetitive cyclic trunk flexion–extension which in turn influences muscular activation including the flexion–relaxation phenomenon. Empirical evidence to support this hypothesis, especially the development of viscoelastic tension–relaxation and its associated muscular response in passive cyclic activity in humans, is incomplete. The objective of this study was to examine the response of lumbar muscles to tension–relaxation development of the viscoelastic tissue during prolonged passive cyclic trunk flexion–extension. Activity of the lumbar muscles remained low and steady during the passive exercise session. Tension supplied by the posterior viscoelastic tissues decreased over time without corresponding changes in muscular activity. Active flexion, following the passive flexion session, elicited significant increase in paraspinal muscles EMG together with increase in the median frequency. It was concluded that reduction of tension in the lumbar viscoelastic tissues of humans occurs during cyclic flexion–extension and is compensated by increased activity of the musculature in order to maintain stability. It was also concluded that the ligamento-muscular reflex is inhibited during passive activities but becomes hyperactive following active cyclic flexion, indicating that moment requirements are the controlling variable. It is conceived that prolonged routine exposure to cyclic flexion minimizes the function of the viscoelastic tissues and places increasing demands on the neuromuscular system which over time may lead to a disorder and possible exposure to injury.     

Decreasing the required lumbar extensor moment induces earlier onset of flexion relaxation

Flexion relaxation (FR) is characterized by the lumbar erector spinae (LES) becoming myoelectrically silent near full trunk flexion. This study was designed to: (1) determine if decreasing the lumbar moment during flexion would induce FR to occur earlier; (2) characterize thoracic and abdominal muscle activity during FR. Ten male participants performed four trunk flexion/extension movement conditions; lumbar moment was altered by attaching 0, 5, 10, or 15 lb counterweights to the torso. Electromyography (EMG) was recorded from eight trunk muscles. Lumbar moment, lumbar flexion and trunk inclination angles were calculated at the critical point of LES inactivation (CPLES). Results demonstrated that counterweights decreased the lumbar moment and lumbar flexion angle at CPLES (p < 0.0001 and p = 0.0029, respectively); the hypothesis that FR occurs earlier when lumbar moment is reduced was accepted. The counterweights did not alter trunk inclination at CPLES (p = 0.1987); this is believed to result from an altered hip to spine flexion ratio when counterweights were attached. Lumbar multifidus demonstrated FR, similar to LES, while thoracic muscles remained active throughout flexion. Abdominal muscles activated at the same instant as CPLES, except in the 15 lb condition where abdominal muscles activated before CPLES resulting in a period of increased co-contraction.     

Active length-tension relation and the effect of muscle pinnation on fiber lengthening

The active length-tension relation was determined for the left digastric muscle of seven New Zealand White rabbits anesthetized with pentobarbital. Measurements of muscle length and fiber architecture were made from photographs of resting and actively contracting muscle. There was a marked difference between length-tension curves based upon resting as compared to active muscle length. The active length-tension relation had a longer descending limb than ascending limb, whereas the length-tension relation based on passive muscle length tended to be symmetrical around optimum length. On the average, muscle fibers lengthened 0.77 mm for each 1 mm of extension of the muscle belly. Since the rabbit digastric muscle is unipinnate, this suggests that pinnation serves to enhance the range of muscle excursion in this muscle.     

Residual force enhancement during voluntary contractions of knee extensors and flexors at short and long muscle lengths

Residual force enhancement (RFE) is a term describing the observation that muscle tension during a contraction that includes a stretch and hold remains above that during an isometric contraction at the hold length. RFE has been observed during in vitro and in vivo experiments, but results involving voluntary contractions are mixed, particularly with respect to large muscles. The purpose of this study was to determine if RFE can be observed in large muscles such as knee extensors and flexors at joint configurations corresponding to the ascending and descending limbs of the muscle force-length curve. Two groups of twenty participants (ten males and ten females per group) performed maximum voluntary contractions on a Biodex machine in purely isometric conditions and in isometric conditions immediately following eccentric stretch. Knee extension trials were performed at 40° (short muscles) and 100° (long muscles) flexion from full extension (0°), and knee flexion trials were performed at 70° (short muscles) and 10° (long muscles) flexion. Stretch-isometric trials terminated at these angles following 30° of eccentric motion at 30°/s. Statistically-significant RFE was observed for both tasks at long-muscle joint configurations, but was not observed for either task at short-muscle joint configurations. Passive torque enhancement was also observed following muscle relaxation at long-muscle joint configurations for both tasks, but for only knee flexion at short-muscle joint configurations. These results reinforce for voluntary contractions of large muscles the RFE behavior observed in smaller muscles, and provide further evidence that RFE occurs primarily on the descending limb of the muscle force-length curve.     

Length-tension relationship of abdominal expiratory muscles: effect of emphysema

The present study examined the active and passive length-tension relationship of the abdominal expiratory muscles in vitro during electrically stimulated contractions. Studies were performed on isolated strips of transverse abdominis and external oblique muscle from nine adult hamsters with normal lung function. The effect of chronic hyperinflation on the two muscles was assessed in eight hamsters with elastase-induced emphysema. In normal animals the maximal active tension per cross-sectional area (Po) was equal in the two muscles. The absolute muscle fiber length at which Po occurred (Lo) was less for the external oblique than the transverse abdominis and the length-tension curve operated at shorter fiber lengths. However, the change in tension produced by an increase or decrease in muscle length expressed in relative terms (i.e., as %Lo) was greater for the transverse abdominis than the external oblique. Mean total lung capacity of emphysematous animals was 198% of control. Po of the transverse abdominis and external oblique were the same in emphysematous and control animals. However, Lo and the length-tension curve of the transverse abdominis occurred at shorter fiber lengths in emphysematous animals because of a reduction in the number of sarcomeres in series along the fiber. The length-tension curve and the number of sarcomeres in the external oblique was the same in emphysematous and control animals. These results in normal animals indicate that the magnitude of the change in active and passive tension produced by a change in muscle length differs in the transverse abdominis and external oblique. Moreover, chronic hyperinflation of the thorax produced by elastase injection alters the length-tension relationships of some but not all the expiratory muscles.     

Ankle flexor muscles in the cat: Length-active tension and muscle unit properties as related to locomotion

The mechanical properties of the whole muscle and fast-twitch muscle units of the cat hindlimb pretibial flexors have been explored and related to normal locomotion. Tibialis anterior (TA) is parallel-fibered and functionally crosses a single joint, the ankle, whereas extensor digitorum longus (EDL) is pinnate and spans the ankle, knee, metatarsophalangeal and interphalangeal joints. The active tetanic tension of TA remains near its peak value over a range of muscle lengths associated with normal ankle movement. In contrast, the length-tension curve of EDL is sharply peaked. However, normal corollary action of the knee, ankle and metatarsophalangeal joints during stepping minimizes EDL's excursion and maintains it at or near a length optimal for peak tension development. EDL is capable of producing synchronous but sterotyped digit and ankle movements while TA provides for independent ankle flexion at all relevant joint angles. The mechanical properties of 84 TA and 98 EDL fast-twitch muscle units were studied by measuring twitch contraction time (≤45 msec), peak tetanic tension, response to repetitive stimulation, and contractile fatigue resistance during electrical stimulation of single alpha axons, functionally isolated from ventral root filaments. These mechanical properties were essentially similar for both muscles with the exception of mean peak tetanic tension which was 30% lower for TA units (14 gm-wt) than for EDL units (20 gm-wt). A high proportion of units in both muscles demonstrated fatigue resistance which is reflective of the repetitive, phasic demand upon these muscles during locomotion.     

Individual muscle force parameters and fiber operating ranges for elbow flexion-extension and forearm pronation-supination

We have quantified individual muscle force and moment contributions to net joint moments and estimated the operating ranges of the individual muscle fibers over the full range of motion for elbow flexion/extension and forearm pronation/supination. A three dimensional computer graphics model was developed in order to estimate individual muscle contributions in each degree of freedom over the full range of motion generated by 17 muscles crossing the elbow and forearm. Optimal fiber length, tendon slack length, and muscle specific tension values were adjusted within the literature range from cadaver studies such that the net isometric joint moments of the model approximated experimental joint moments within one standard deviation. Analysis of the model revealed that the muscles operate on varying portions of the ascending limb, plateau region, and descending limb of the force-length curve. This model can be used to further understand isometric force and moment contributions of individual muscles to net joint moments of the arm and forearm and can serve as a comprehensive reference for the forces and moments generated by 17 major muscles crossing the elbow and wrist.     

The effect of rate, paired pacing and calcium on the length-tension relations in the rabbit papillary muscle

The effect of sustained paired pacing, extracellular Ca2+ concentration and of the rate of steady-state single pacing on the length-tension relations in the isolated papillary muscles of right ventricles of rabbit hearts was investigated. We found that full range change of contractile force (CF, from close to zero to maximal) may be obtained by means of the change of the resting muscle length by 10-15% of Lmax (the length at which CF is maximal). This change brings about the shift of the thin filaments of the sarcomeres along the thick ones by one sixth of their length. The only effect of all the applied interventions could be reproduced by multiplying CF at each length step by some coefficient. Neither of them did change the basic pattern of length-tension relations. These results are compatible with the hypothesis that the basic mechanism of length-tension relations is the change of sensitivity of contractile system to the sarcoplasmic Ca2+ concentration.     

Relationship between length-tension relation and 20,000 dalton myosin light chain phosphorylation in guinea-pig taenia caeci

1. Relationship between length-tension relation and phosphorylation of 20,000 dalton myosin light chain (LC20) in guinea-pig taenia caeci was investigated. 2. At in situ length (Lb), a good linear correlation was obtained between isometric tension and LC20 phosphorylation in high-K+-stimulated muscle. 3. In 100 mM K+-stimulated muscle, the active tension decreased at muscle lengths other than Lb, but no significant decrease in degree of LC20 phosphorylation was observed. 4. These results suggest that in guinea-pig taenia caeci, the major portion of the decrease in active tension at muscle lengths other than Lb is not due to a decrease in degree of activation.     

Muscular contributions to hip and knee extension during the single limb stance phase of normal gait: a framework for investigating the causes of crouch gait

Crouch gait, a troublesome movement abnormality among persons with cerebral palsy, is characterized by excessive flexion of the hips and knees during stance. Treatment of crouch gait is challenging, at present, because the factors that contribute to hip and knee extension during normal gait are not well understood, and because the potential of individual muscles to produce flexion or extension of the joints during stance is unknown. This study analyzed a three-dimensional, muscle-actuated dynamic simulation of walking to quantify the angular accelerations of the hip and knee induced by muscles during normal gait, and to rank the potential of the muscles to alter motions of these joints. Examination of the muscle actions during single limb stance showed that the gluteus maximus, vasti, and soleus make substantial contributions to hip and knee extension during normal gait. Per unit force, the gluteus maximus had greater potential than the vasti to accelerate the knee toward extension. These data suggest that weak hip extensors, knee extensors, or ankle plantar flexors may contribute to crouch gait, and strengthening these muscles--particularly gluteus maximus--may improve hip and knee extension. Abnormal forces generated by the iliopsoas or adductors may also contribute to crouch gait, as our analysis showed that these muscles have the potential to accelerate the hip and knee toward flexion. This work emphasizes the need to consider how muscular forces contribute to multijoint movements when attempting to identify the causes of abnormal gait.     

Length-tension relationships are altered in regenerating muscles of the rat after bupivacaine injection.

Intramuscular injection of bupivacaine causes complete degeneration of fibers in extensor digitorum longus (EDL) muscles of rats, followed by complete regeneration within 60 days. Previous studies have shown that regenerated EDL muscles are protected from contraction-induced injury 60 days after bupivacaine injection. It is possible that these regenerated muscles have altered length-tension relations because of fiber remodeling. We tested the hypothesis that length-tension relations are different in bupivacaine-injected and noninjected control muscles. EDL and soleus muscles of the right hindlimb of deeply anesthetized rats were injected with bupivacaine and then allowed to recover for 7, 14, 21, or 60 days (7D, 14D, 21D, 60D), and isometric contractile properties were assessed. Muscles of the contralateral limb were not injected and served as control. EDL muscles recovered from bupivacaine injection more rapidly than soleus muscles, with mass restored to control levels at 21D, and isometric tetanic force (P(o)) restored to control at 60D. In contrast, mass and P(o) of injected soleus muscles was not restored to control even at 60D. In 7D EDL muscles, length-tension curves were shifted leftward compared with control, but in 21D and 60D EDL muscles length-tension curves were right shifted significantly (treatment x muscle length: P < 0.001). Although no clear shift in the position of the length-tension curve was observed in regenerating soleus muscles, force production was enhanced on the descending limb of the curve in 60D soleus muscles (treatment x relative muscle length: P < 0.01). The rightward shift in the length-tension curve of EDL muscles 60 days after bupivacaine injection is likely to contribute to the mechanism for their previously observed protection from contraction-induced injury.     

Rises in whole muscle passive tension of mammalian muscle after eccentric contractions at different lengths.

This is a report of experiments carried out on the medial gastrocnemius muscle of the anesthetized cat, investigating the effects of eccentric contractions carried out at different muscle lengths on the passive and active length-tension relationships. In one series of experiments, the motor supply to the muscle was divided into three approximately equal parts; in the other, whole muscles were used. Fifty eccentric contractions were carried out over different regions of the active length-tension curve for each partial or whole muscle. Active and passive length-tension curves were measured before and after the eccentric contractions. When eccentric contractions were carried out at longer lengths, there was a larger shift of the optimum length for active tension in the direction of longer muscle lengths and a larger fall in peak isometric tension. Passive tension was higher immediately after the eccentric contractions, and if the muscle was left undisturbed for 40 min, it increased further to higher values, particularly after contractions at longer lengths. A series of 20 passive stretches of the same speed and amplitude and covering the same length range as the active stretches, reduced the passive tension which redeveloped over a subsequent 40-min period. It is hypothesized that there are two factors influencing the level of passive tension in a muscle after a series of eccentric contractions. One is injury contractures in damaged muscle fibers tending to raise passive tension; the other is the presence of disrupted sarcomeres in series with still-functioning sarcomeres tending to reduce it.     

Links between the mechanics of ventilation and spine stability

Spine stability is ensured through isometric coactivation of the torso muscles; however, these same muscles are used cyclically to assist ventilation. Our objective was to investigate this apparent paradoxical role (isometric contraction for stability or rhythmic contraction for ventilation) of some selected torso muscles that are involved in both ventilation and support of the spine. Eight, asymptomatic, male subjects provided data on low back moments, motion, muscle activation, and hand force. These data were input to an anatomically detailed, biologically driven model from which spine load and a lumbar spine stability index was obtained. Results revealed that subjects entrained their torso stabilization muscles to breathe during demanding ventilation tasks. Increases in lung volume and back extensor muscle activation coincided with increases in spine stability, whereas declines in spine stability were observed during periods of low lung inflation volume and simultaneously low levels of torso muscle activation. As a case study, aberrant ventilation motor patterns (poor muscle entrainment), seen in one subject, compromised spine stability. Those interested in rehabilitation of patients with lung compromise and concomitant back troubles would be assisted with knowledge of the mechanical links between ventilation during tasks that impose spine loading.     

Relation between the ankle joint angle and the maximum isometric force of the toe flexor muscles

The purpose of this study was to investigate the relationships between the ankle joint angle and maximum isometric force of the toe flexor muscles. Toe flexor strength and electromyography activity of the foot muscles were measured in 12 healthy men at 6 different ankle joint angles with the knee joint at 90 deg in the sitting position. To measure the maximum isometric force of the toe flexor muscles, subjects exerted maximum force on a toe grip dynamometer while the activity levels of the intrinsic and extrinsic plantar muscles were measured. The relation between ankle joint angle and maximum isometric force of the toe flexor muscles was determined, and the isometric force exhibited a peak when the ankle joint was at 70–90 deg on average. From this optimal neutral position, the isometric force gradually decreased and reached its nadir in the plantar flexion position (i.e., 120 deg). The EMG activity of the abductor hallucis (intrinsic plantar muscle) and peroneus longus (extrinsic plantar muscle) did not differ at any ankle joint angles. The results of this study suggest that the force generation of toe flexor muscles is regulated at the ankle joint and that changes in the length-tension relations of the extrinsic plantar muscle could be a reason for the force-generating capacity at the metatarsophalangeal joint when the ankle joint angle is changed.     

Length-tension relation in Limulus striated muscle

Laser diffraction techniques coupled with simultaneous tension measurements were used to determine the length-tension relation in intact, small (0.5-mm thick, 10-mm wide, 20-25-mm long) bundles of a Limulus (horseshoe crab) striated muscle, the telson levator muscle. This muscle differs from the model vertebrate systems in that the thick filaments are not of a constant length, but shorten from 4.9 to approximately 2.0 micrometers as the sarcomeres shorten from 7 to 3 micrometers. In the Limulus muscle, the length-tension relation plateaued to an average maximum tension of 0.34 N/mm2 at a sarcomere length of 6.5 micrometers (Lo) to 8.0 micrometers. In the sarcomere length range from 3.8 to 12.5 micrometers, the muscle developed 50% or more of the maximum tension. When the sarcomere lengths are normalized (expressed as L/Lo) and the Limulus data are compared to those from frog muscle, it is apparent that Limulus muscle develops tension over a relatively greater range of sarcomere lengths.