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Osmotic compression and stiffness changes in relaxed skinned cardiac myocytes in PVP-40 and dextran T-500.

Sarcomere lengths, cell widths, indices of stiffness, and striation pattern uniformity were determined from radially compressed isolated adult cardiac myocytes from the rat. Single cells were bathed in a series of relaxing solutions containing 0-15% concentrations of nonpenetrating long chain polymers PVP-40 and dextran T-500. There were no significant changes observed in average sarcomere lengths or in striation pattern uniformity at any concentration. But cell widths decreased and stiffness increased in both polymers in a concentration-osmotic pressure-dependent relationship. Changes in cell width and stiffness were repeatable in either polymer, but only after an initial compression with a 10 or 15% concentration solution. The observed reduction in cell width after initial compression correlates well with known myofilament lattice spacing compression in rat cardiac muscle and is qualitatively similar to compressions seen in skeletal muscle preparations. But the cardiac myofilament lattice may not be as compressible as the skeletal lattice. Like skeletal muscle, stiffness exhibits a two-phase relationship where most of the increase occurs at solution osmotic pressures greater than 20 Torr. Finally, the inherently greater passive stiffness-length relationship of cardiac muscle is maintained at higher osmotic pressures such that the passive elastic modulus is strongly length dependent.

Actin Cytoskeleton↗

Muscle and tendon stiffness in patients with upper motor neuron lesion following a stroke.

The objective of this study was to investigate muscle and tendon stiffness in the triceps surae muscles in patients who had previously had a stroke. The participants were 12 men showing slight to moderate degrees of muscle tonus in the affected leg. All patients showed minimal or no overt clinical motor symptoms, and all walked without mechanical aid. Muscle strengths in isometric and isokinetic activities were measured, as was passive resistance during plantarflexion in each leg. Walking speed was also measured. Evaluations of physical performance and muscle tone were made. Muscle and tendon stiffness was calculated from measurements whilst passively stretching during electrical stimulation, separately for each leg. Muscle strength was significantly higher in the non-affected than in the affected leg. Muscle stiffness was significantly higher in the affected leg than in the non-affected leg. Tendon stiffness was significantly higher in the non-affected than in the affected leg. The higher muscle stiffness in the affected leg might enhance the possibility for storing elastic energy during preactivation. Lower tendon stiffness in the affected leg might reduce the development of fatigue in movements at low velocities.

Adult↗

Magnetic resonance elastography of skeletal muscle.

While the contractile properties of skeletal muscle have been studied extensively, relatively little is known about the elastic properties of muscle in vivo. Magnetic resonance elastography (MRE) is a phase contrast-based method for observing shear waves propagating in a material to determine its stiffness. In this work, MRE is applied to skeletal muscle under load to quantify the change in stiffness with loading. A mathematical model of muscle is developed that predicts a linear relationship between shear stiffness and muscle load. The MRE technique was applied to bovine muscle specimens (N = 10) and human biceps brachii in vivo (N = 5). Muscle stiffness increased linearly for both passive tension (14.5 +/- 1.77 kPa/kg) and active tension, in which the increase in stiffness was dependent upon muscle size, as predicted by the model. A means of noninvasively assessing the viscoelastic pro-perties of skeletal muscle in vivo may provide a useful method for studying muscle biomechanics in health and disease.

Animals↗

Relationship between muscle force and stiffness in the whole mammalian muscle: a simulation study.

Several types of analyses in biomechanics require estimates of both muscle force and stiffness. Simulations were performed using the two-state cross-bridge Bond Distribution-Moment muscle model of Zahalak (1981), together with other parameters for passive elasticity and tendon compliance, to estimate instantaneous stiffness and to compare these estimates with the wide range of values reported in the literature. While the relatively simple cross-bridge theory appears to approximate the stiffness of skinned muscle fibers, the stiffness of a complete muscle-tendon unit become complex and non-linear due to relative changes in muscle-tendon length and interaction with activation and length dependent passive elastic components. It would appear that the variability in muscle stiffness values reported in the literature can be explained with the D-M approach.

Biomechanical Phenomena↗

Desmin integrates the three-dimensional mechanical properties of muscles.

Striated muscle is a linear motor whose properties have been defined in terms of uniaxial structures. The question addressed here is what contribution is made to the properties of this motor by extramyofilament cytoskeletal structures that are not aligned in parallel with the myofilaments. This question arose from observations that transverse loads increase muscle force production in diaphragm but not in the hindlimb muscle, thereby indicating the presence of structures that couple longitudinal and transverse properties of diaphragmatic muscle. Furthermore, we find that the diaphragms of null mutants for the cytoskeletal protein desmin show 1) significant reductions in coupling between the longitudinal and transverse properties, indicating for the first time a role for a specific protein in integrating the three-dimensional mechanical properties of muscle, 2) significant reductions in the stiffness and viscoelasticity of muscle, and 3) significant increases in tetanic force production. Thus desmin serves a complex mechanical function in diaphragm muscle by contributing both to passive stiffness and viscoelasticity and to modulation of active force production in a three-dimensional structural network. Our finding changes the paradigm of force transmission among cells by placing our understanding of the function of the cytoskeleton in the context of the structural and mechanical complexity of muscles.

Animals↗

The effect of aging on ventricular contractile performance.

The effects of aging on mechanical performance of isolated canine right ventricular trabeculae were studied in two age groups. The first group was comprised of nine dogs, about 9 months of age. The second group was composed of seven dogs over 8 years of age. Aging had no significant effect on developed force. Extent of shortening tended to decrease. There was a significant decrease in both the rate of rise of tension and the velocity of shortening (20%). THis reduction was primarily due to an increased duration of contraction. Twitch duration increased by as much as 40% during aging but most of this prolongation was due to a profound slowing of relaxation. Aging caused a significant increase in passive stiffness since equivalent changes in muscle length brought about twice as much increase in resting tension in the aged muscle as in the young muscle. On the other hand, aging caused a significant shift of Lo to the right. Taken collectively, these results indicate that aging is associated with increased passive stiffness and decreased speed of contraction without changes in strength.

Aging↗

Changes in arterial wall properties during development and maintenance of renal hypertension.

Segments of carotid and tail arteries were used to determine the effects of two-kidney Goldblatt hypertension on mechanical properties and chemical contents. Arteries were obtained after 4 and 12 wk of hypertension. Systolic pressures averaged 129 mmHg in control animals and 150 mmHg in hypertensive ones. Pressure-diameter data were recorded under conditions of active (145 mM K+) and passive (0 Ca2+ and 2 mM EGTA) smooth muscle. Small increases in passive stiffness were found after 4 and 12 wk in the carotid arteries of hypertensive animals, but no significant change occurred in the tail arteries. Small but significant decreases in total connective tissue content of these arteries were found after 12 wk of hypertension. Maximum values of active stress response were significantly larger in carotid arteries from hypertensive animals at both 4 and 12 wk of hypertension. No significant differences were found for the tail arteries. The K+ content of the carotid but not the tail artery was larger for the hypertensive animals. When corrected for differences in relative cell content most of the differences in active force development were eliminated. The differences in passive mechanics can not be explained on the basis of the changes in connective tissue content. The active smooth muscle responses and content changes suggest that they are directly related to the moderate elevation of arterial pressure.

Animals↗

Biomechanical responses to repeated stretches in human hamstring muscle in vivo.

To examine stiffness, energy, and passive torque in the dynamic and static phases of a stretch maneuver in the human hamstring muscle in vivo we used a test-retest protocol and a repeated stretches protocol. Resistance to stretch was defined as passive torque (in newton-meters) offered by the hamstring muscle group during passive knee extension as measured using an isokinetic dynamometer with a modified thigh pad. In 13 uninjured subjects, the knee was passively extended to a predetermined final position (0.0875 rad/ sec, dynamic phase) where it remained stationary for 90 seconds (static phase). The test-retest protocol included two tests administered 1 hour apart. On a separate occasion, five consecutive static stretches were administered separated by 30 seconds and followed by a sixth stretch 1 hour later. For the test-retest phase, stiffness and energy in the dynamic phase and passive torque in the static phase did not differ and yielded correlations of r = 0.91 to 0.99. During the static phase, passive torque declined in both tests (P < 0.0001). For the repeated stretches, decreases were observed for energy (P < 0.01) and stiffness (P < 0.05) in the dynamic phase and for passive torque (P < 0.0001) in the static phase. However, the decline in the variables returned to baseline within 1 hour. The data show that the method employed is a useful tool for measuring biomechanical variables during a stretch maneuver. This may provide a more detailed method to examine skeletal muscle flexibility.

Adult↗

Efficiency of work production by spastic muscles.

The present study compared the muscular efficiency in spastic and healthy lower limbs producing the same mechanical work. Sixteen chronic post-stroke hemiparetic and spastic patients and 14 age-matched healthy subjects were submitted to a submaximal stepwise exercise testing on a bicycle ergometer, pedalling with only one lower limb. Net energetic expenditure was computed from oxygen consumption above resting values. Electrical activity of antagonistic muscles in the thigh and in the shank was recorded and co-contraction was defined as the percentage of the pedalling cycle when antagonistic muscles were activated simultaneously. The efficiency was calculated as the ratio between the mechanical work done on the ergometer and the net energetic expenditure. Spasticity was quantitatively evaluated by measuring passive ankle plantar flexor muscle stiffness. The working capacity of the patients' paretic lower limb was very low (<40W). The energy expenditure increased linearly as a function of work intensity, without statistical difference between the patients paretic lower limb (PPL), the patients healthy lower limb (PHL) and the healthy subjects lower limb (HSL). Shank co-contraction was 2.9 times greater in PPL (p<0.05) and 2.3 times greater in PHL (p<0.05) than in HSL. Thigh co-contraction was also 1.8 times greater in PPL than in HSL (p<0.05). The ankle plantar flexor muscle stiffness was statistically greater in PPL than in PHL and HSL (p<0.05). The efficiency was not statistically different between the three groups (p=0.155). In conclusion, the efficiency of work production by paretic and spastic lower limb muscles was normal ( congruent with 20%) despite significant neurological impairments.

Adult↗

Multi-axis passive and active stiffnesses of the glenohumeral joint.

OBJECTIVE: To investigate passive and active glenohumeral stiffness in the anterior, posterior, superior, and inferior directions at different lateral positions of the humerus. DESIGN: Glenohumeral stiffness along multiple axes was determined in fresh-frozen shoulder specimens under both passive (no simulated muscle contraction) and active (with simulated muscle contraction) conditions. BACKGROUND: Glenohumeral laxity has been evaluated in various studies with focus on one of the multiple directions. However, glenohumeral stiffness characterizing the force-displacement relationship and stability has not been evaluated in all four directions under passive and active conditions. METHODS: The humeral head was translated in the posterior, anterior, inferior and superior directions relative to the glenoid with different lateral positions, and multi-axis glenohumeral stiffness generated by passive and active structures were investigated. RESULTS: Without muscle loading, glenohumeral stiffness in the superior direction ( Ksup = 5.83 N/mm) was higher than that in the inferior ( Kinf = 4.32 ), anterior ( Kant = 3.67 ), and posterior ( Kpost = 2.89 ) directions ( P < 0.008 ), and Kinf was higher than Kpost ( P = 0.011 ). Stiffness in the different directions were correlated to each other ( P < 0.001 ), and shifting the humerus laterally increased stiffness in all directions ( P < 0.05 ) except for the superior direction. With moderate muscle loads, the glenohumeral joint became significantly stiffer in all four directions ( P < 0.05 ) with less difference among different directions. CONCLUSIONS: Glenohumeral stiffnesses are different in the different directions but are correlated to each other and contribute jointly to glenohumeral stability. Muscle contractions can increase glenohumeral stiffness significantly.

Biomechanical Phenomena↗

Connective tissue changes and physical properties of developing and ageing skeletal muscle.

Selective staining of the connective tissue and image analysis showed that in the extensor digitorum longus and soleus muscles of the rat there was an increase in the thickness of the endomysium in both early growth and senility. The perimysium thickness was more or less constant throughout life except in senility when the concentration of this component also increased. The stiffness (length-passive tension) of these muscles was found to increase throughout life. Log transforms of the length-passive tension plots had particularly steep slopes in the senile extensor digitorum longus muscle. Except in the senile soleus muscle, the increase in stiffness was closely correlated with the increase in endomysium and perimysium and with total muscle collagen (as measured biochemically) with age. The relationship between the initial length and the active tension in the extensor digitorum longus muscle changed with age. The older muscles showed a greater decline in tension for each decrement of length resulting from the increased development of the connective tissue.

Aging↗

Do cross-bridges contribute to the tension during stretch of passive muscle?

The tension rise during stretch of passive skeletal muscle is biphasic, with an initial steep rise, followed by a subsequent more gradual change. The initial rise has been interpreted as being due to the presence of numbers of long-term, stable cross-bridges in resting muscle fibres. A point of weakness with the cross-bridge interpretation is that the initial stiffness reaches its peak value at muscle lengths beyond the optimum for myofilament overlap. To explain this result it has been suggested that despite the reduced overlap at longer lengths, the closer interfilament spacing and a higher sensitivity of the myofilaments to Ca2+ allows more stable cross-bridges to form. Recently the stretch responses of passive muscle have been re-examined and it has been suggested that it is not necessary to invoke cross-bridge mechanisms at all. Explanations based on a viscous resistance to interfilament sliding and mechanical properties of the elastic filaments, the gap filaments, were thought to adequately account for the observed tension changes. However, an important property of passive muscle, the dependence of stretch responses on the immediate history of contraction and length changes, thixotropy, cannot be explained simply in terms of viscous and viscoelastic properties. The review discusses the cross-bridge interpretation of muscle thixotropy and the relationship of passive stiffness to filament resting tension and latency relaxation. It is proposed that cross-bridges can exist in three states; one, responsible for the resting stiffness, requires resting levels of calcium. When, during activation, calcium levels rise, cross-bridges enter a low-force, high-stiffness state, signalled by latency relaxation, before they move to the third, force-generating state. It is concluded that, compared with viscoelastic models, a cross-bridge-based explanation of passive muscle properties is better able to accommodate the currently known facts although, as new information becomes available, this view may need to be revised.

Actin Cytoskeleton↗

Evaluation of triceps surae muscle length and resistance to passive lengthening in patients with acquired brain injury.

OBJECTIVE: To examine changes in muscle length and resistance to passive lengthening in the triceps surae muscles in patients with recently acquired brain injury. BACKGROUND: Increased passive resistance in the triceps surae muscles is common following acquired brain injury. Adaptive shortening secondary to relative immobility, and increased stiffness due to rheologic changes within the musculo-tendinous unit, may be exacerbated by plantarflexor muscle overactivity related to the brain injury itself. DESIGN: Three variables representing resistance to passive lengthening and soleus muscle length were compared between subjects with recent brain injury and age matched normal controls. Comparison between limbs was made for subjects with unilateral neurological impairment. METHODS: Slow passive dorsiflexion stretches were performed using a computer controlled dynamometer. Muscle stiffness in the initial and latter portion of the range, and the angles achieved at torques of 5 and 10 N m were determined from torque-angle curves. Maximal ankle dorsiflexion with the knee flexed was considered to reflect soleus muscle length. RESULTS: Significant differences were demonstrated for all variables, except passive stiffness near the end of available range. The limb ipsilateral to unilateral brain injury differed from control limbs in that significantly less passive range of dorsiflexion was available and initial resistance to passive stretch was significantly less. CONCLUSIONS: The reduction in soleus muscle length evident in subjects with recent acquired brain injury, even in neurologically unaffected limbs, may reflect the influence of relative immobility. Although plantarflexor muscle overactivity was found to be associated with increased resistance to slow passive stretch, the mechanism was unable to be elucidated from these data. The limb ipsilateral to unilateral neurological impairment cannot be considered to be a 'normal' control for comparative purposes. RELEVANCE: Adaptive shortening and increased resistance to passive lengthening limit active ankle dorsiflexion, and alter ankle biomechanics. Tonic muscle overactivity has the potential to exacerbate these changes. Prophylactic management of inappropriate muscle activity and maintenance of muscle length may facilitate the achievement of rehabilitation goals and reduce subsequent disability following acquired brain injury.

Adolescent↗

Stretching exercises: effect on passive extensibility and stiffness in short hamstrings of healthy subjects.

Passive muscle stretch tests are common practice in physical therapy and rehabilitation medicine. However, the effects of stretching exercises are not well known. With an instrumental straight-leg-raising set-up the extensibility, stiffness, and electromyographic activity of the hamstring muscles have been experimentally determined and the effects of stretching exercises have been evaluated. Fourteen volunteers, aged 20 to 38 years (mean 27.3) were selected from a young healthy population with the toe-touch test (finger-ground distance greater than 0cm), and a straight-leg-raising angle about 80 degrees. According to usual standards the diagnosis was short hamstrings. One group of seven subjects was treated during 4 weeks with a daily home exercise program aimed at stretching the hamstrings, whereas the untreated group was used as a control. Instrumental straight-leg-raising was performed in the subjects of both groups. The significance of the differences between the mean values was determined with the Student's t-test. Comparison of the data obtained before and after the muscle stretching program showed a slight but significant increase in the extensibility of the hamstrings accompanied with a significant increase of the stretching moment tolerated by the passive hamstring muscles. However, the elasticity remained the same. It is concluded that stretching exercises do not make short hamstrings any longer or less stiff, but only influence the stretch tolerance.

Adult↗

Biaxial constitutive relations for the passive canine diaphragm.

Samples of the muscular sheet excised from the midcostal region of dog diaphragms were subjected to biaxial loading. That is, stresses in the direction of the muscle fibers and in the direction perpendicular to the fibers in the plane of the sheet were measured at different combinations of strains in the two directions. Stress-strain relations were obtained by fitting equations to these data. In the direction of the muscle fibers, for strains up to 0.7, stress is a modestly nonlinear function of strain and ranges up to approximately 60 g/cm. In the direction perpendicular to the fibers, the sheet is stiffer and more strongly nonlinear. At a strain in the perpendicular direction of approximately 0.35, stress increases abruptly. The stress-strain relation in the muscle direction is consistent with observations of passive muscle shortening in vivo. However, the stiffness in the perpendicular direction is not high enough to explain the observation that strains in the perpendicular direction in vivo are nearly zero. We conclude that, in the passive diaphragm in vivo, stress in the direction perpendicular to the muscle fibers is small.

Animals↗

Effects of tetanic contraction of motor units of similar type on the initial stiffness to ramp stretch of the cat peroneus longus muscle.

1. The stiffness during the initial portion of a ramp stretch was measured in cat peroneus longus muscle at rest and during maximal tetanic contractions produced by increasing numbers of motor units of the same type [slow (S), fast fatigue resistant (FR), or fast fatigable (FF)]. 2. This initial ramp stiffness was defined as the ratio between tension and length change over the limited range of constant velocity extension during which tension rose linearly with length change. This stiffness was reduced by tetanic contraction of a number of motor units while other units remained inactive. The reduction had different characteristics in contractions produced by S, FR, or FF units. 3. Two brief ramp (triangular) stretches were applied at short intervals to evaluate the contribution of stable cross bridges to the changes in ramp stiffness. When the amplitude of the first stretch exceeded the presumed elastic limits of the stable cross bridges, the second ramp stretch showed a reduction of 20-60% in initial stiffness. This was seen both in passive muscles and in muscles in which several motor units were contracting. 4. When increasing numbers of motor units of the same type were activated, the initial ramp stiffness to the second of a pair of triangular stretches delivered during contraction increased almost linearly with the developed tension. The slope of this increase was 2.5 times steeper for S units than for FR units. This reflects the fact that contraction produced by S units causes a proportionally greater resistance to stretch than that of fast units.

Animals↗

Mechanical behaviour of hamstring muscles in low-back pain patients and control subjects.

The purpose of this study was to measure and compare the passive elastic moment, the stiffness and the damping coefficient of the hip joint, as functions of the hip and knee joint angles in men with and without low-back pain. Two conventional tests, the straight-leg-raising test and the trunk forward flexion, were also performed and compared between these subjects. The passive elastic moment was measured using an isokinetic device in the passive mode. This device raised the lower limb from the horizontal position to the straight-leg-raising angle at a slow and constant angular velocity. A custom-made splint connected with the lever arm of the isokinetic device maintained the knee in extension and the ankle in the neutral position. The damping coefficient of the hip joint was measured for 0, 15, 45, 60, 75 and 90% of straight leg raising angle of each subject, using the suspension method based on small oscillation theory. To ensure that muscles were inactive during the passive hip moment tests, muscle activity was monitored with surface EMG. The stiffness was computed as the ratio of the change in passive elastic moment to the change in the hip angle. The passive elastic moment, the stiffness and the normalized trunk flexion were significantly different between the two groups respectively. There was, however, no difference between the two groups in the results of straight-leg-raise and damping coefficient of the hip. The passive elastic moment was a nonlinear function of the hip flexion angle and showed large intersubject differences, especially as the joint limit was approached. The damping coefficient was a polynomial function of the hip flexion angle. The measured variables were analysed using a discriminant function and it was shown that the two groups were clearly discriminable in a meaningful manner.

Journal Article↗

The effects of neuromuscular stimulation on muscle tone at the knee in paraplegia.

Resting muscle tone of the leg was measured in terms of thigh muscle stiffness and knee resonant frequency in muscles of spinal cord injured subjects who had been involved in an electrical neuromuscular stimulation training programme of the thigh muscles over at least 2 months. The thigh circumference of these patients was 6.6% larger than before training commenced (P less than 0.001) and showed increased muscle stiffness and resonant frequency compared to a similar group of paralysed subjects who had not used any neuromuscular stimulation. Resonant frequency and stiffness after the long-term training were similar to those of non-injured controls and therefore the stimulation programme seemed to reverse the effects of paralysis on muscle tone. Short periods of rest (30 min) caused increased muscle stiffness in non-injured controls and paralysed muscles trained by neuromuscular stimulation. Additional 15 min periods of neuromuscular stimulation further increased muscle stiffness in the trained muscles but also in the muscles of paralysed subjects who had no long-term neuromuscular training. In contrast, 15 min sessions of passive movement of the knee decreased muscle stiffness in long-term trained paralysed muscles and untrained paralysed muscles. Knee resonant frequency was also significantly decreased in the trained paralysed muscles. Results show that muscle tone varies depending on the amount of previous movement or rest and that although neuromuscular stimulation of paralysed muscles increases muscle stiffness and knee resonant frequency, it is in fact restoring such properties of the muscle to a state approaching that of non-injured controls.

Adult↗