A quantitative model of intersarcomere dynamics during fixed-end contractions of single frog muscle fibers.

A numerical model of a muscle fiber as 400 sarcomeres, identical except for their initial lengths, was used to simulate fixed-end tetanic contractions of frog single fibers at sarcomere lengths above the optimum. The sarcomeres were represented by a lumped model, constructed from the passive and active sarcomere length-tension curves, the force-velocity curve, and the observed active elasticity of a single frog muscle fiber. An intersarcomere force was included to prevent large disparities in lengths of neighboring sarcomeres. The model duplicated the fast rise, slow creep rise, peak, and slow decline of tension seen in tetanic contractions of stretched living fibers. Decreasing the initial non-uniformity of sarcomere length reduced the rate of rise of tension during the creep phase, but did not decrease the peak tension reached. Limitations of the model, and other processes that might contribute to the shape of the fixed end tetanic tension record are discussed. Taking account of model and experimental results, it is concluded that the distinctive features of the tension records of fixed end tetanic contraction at lengths beyond optimum can be explained by internal motion within the fiber.     

Spatial and temporal heterogeneity of superficial muscle strain during in situ fixed-end contractions

Numerical models of contracting muscle offer a powerful tool to study local mechanical load. For validation of these models, the spatial and temporal distribution of strain was quantified in fixed-end contracting rat tibialis anterior muscle in situ at optimal muscle length (L(o)) and at 120 degrees plantar flexion as well as at 125 and 33Hz stimulation frequency. We studied the hypothesis that after termination of stimulation in situ muscle segments near the motor endplates elongate while segments away from the endplates shorten. We show that both spatial and temporal inhomogeneities in muscle deformation occurred during contraction. Muscle plateau shortening strain equalled 4.1%. Maximal plateau shortening of a muscle segment was much larger (9.6%) and occurred distally (at 0.26 of the scaled length of the muscle). Manipulating torque levels by decreasing the stimulation frequency at the same muscle length induced a decrease in torque ( approximately 20%) with a smaller effect on the level and no effect on the pattern of muscle deformation. During relaxation, distal segments actively shortened at the expense of proximal muscle segments, which elongated. The segments undergoing lengthening were nearer to motor endplates than segments undergoing shortening.In conclusion, the present study provides experimental data on magnitude of contraction-induced deformation needed for validation of numerical models. Local muscle deformation is heterogeneous both temporally and spatially and may be related to proximity to the motor endplates.     

Medial gastrocnemius muscle-tendon interaction and architecture change during exhaustive hopping exercise

Background: Previous literature has shown in vivo changes in muscle-tendon interaction during exhaustive stretch-shortening cycle (SSC) exercise. It is unclear whether these changes in muscle-tendon length during exhaustive SSC exercise are associated with changes in mechanical efficiency (ME). The purpose of the study was to investigate whether changes in platarflexor contractile component (CC) length, tendon length, and changes in plantarflexor muscle activity could explain reduction in ME during exhaustive SSC exercise. Methods: Eight males participated in an exhaustive hopping task to fatigue. Mechanical work and energy expenditure were calculated at different time-points during the hopping task. Furthermore, hopping kinetics and kinematics, medial gastrocnemius (MG) muscle activity, and in vivo ultrasound of the MG were also collected at different time-points throughout the hopping task. Results: ME did not change during the hopping protocol despite shorter tendon and longer CC lengths as subjects approached exhaustion. Percent decreases in pennation angle and muscle thickness were most strongly correlated to time to exhaustion (r = 0.94, p ⩽ 0.05; r = 0.87, p ⩽ 0.05; respectively). Percent changes in CC length change and pennation angle were strongly correlated to percent decrease in maximal voluntary isometric plantarflexion (MVIP) force (r = −0.71, p ⩽ 0.04; r = 0.70, p ⩽ 0.05; respectively). Braking/push-off EMG ratio increased from initial pre-fatigue values to all other time points showing neuromuscular adaptations to altered muscle lengths. Conclusion: Findings from the current study suggest that changes in CC and tendon lengths occur during repetitive hopping to exhaustion, with the amount change strongly related to time to exhaustion. ME of hopping remained unchanged in the presence of altered CC and tendon lengths.     

Nonsteady motion in unloaded contractions of single frog cardiac cells.

We studied the mode of shortening of enzymatically isolated single frog cardiac cells with a high-speed videosystem to see whether or not shortening is smooth. The segmental shortening of the cell in response to electrical stimulation exhibited a clear pause following the initial shortening over a distance of approximately 11 nm/half-sarcomere. Several preparations showed a second pause following the initial one. Nonsteady motion with a pause lasted usually a few tens of milliseconds. The duration of nonsteady motion was shorter in cells with large velocities of steady shortening following the pause than those with smaller velocities.     

Model of the electromyographic activity during brief isometric contractions

EMG's of short isometric contractions are analysed. Mathematical expressions are derived for the average EMG and for the variance of the EMG. The result shows that by deconvolution methods a distribution of motor unit onset time during contraction can be obtained once the mean motor unit potential is known. A further result of the analysis is the fact that by simulation methods a statistic can be derived to estimate the mean number of active motor units covered by the electrodes. The fact that highly variable EMG's go along with very accurately timed movements is shortly discussed.     

Intersarcomere dynamics of single muscle fibers during fixed-end tetani

The contraction dynamics of end and center regions of single fibers have been measured during fixed-end tetani. Experimental control and data acquisition are provided by a digital system that can acquire diffraction data as fast as every 260 microseconds for 300-700 ms. Tension records are simultaneously displayed on a storage oscilloscope. Resting sarcomere length variation between the end and center regions was analogous to that of Gordon et al. (1966). During the rapid rise in force (less than 45 ms), the end regions contract almost twice as fast as the center regions. During the slow rise in force, the velocity of contraction of the end regions was 3.8 times the velocity of stretch of the center regions. In addition, factors that affected the rate and extent of the slow rise in tension also affected the rate and extent of end shortening. In 58% of the cases studied, the amount of shortening observed in the end region was enough to explain the extent of the slow rise in tension. These data support the explanation of creep first proposed by A. V. Hill (1953) and used by Gordon et al. (1966) to justify their use of the back-extrapolation technique in measuring the isometric force-generating capability of a single fiber. These data also indicate that the laser diffraction technique may provide an effective, noninvasive method for studying sarcomere dynamics during creep and related phenomena.     

Energy liberation and chemical change in frog skeletal muscle during single isometric tetanic contractions

Recent data obtained from Rana temporaria sartorius muscles during an isometric tetanus indicate that the time-course of phosphocreatine (PC) splitting cannot account for the total energy (heat + work) liberation (Gilbert et al. 1971. J. Physiol. (Lond.) 218:)63). As this conclusion is important to an understanding of the chemical energetics of contraction, similar experments were performed on unpoisoned, oxygenated Rana pipiens sartorius muscles. The muscles were tetanized (isometrically) at 0 degrees C for 0.6, 1, or 5 s; metabolism was rapidly arrested by freezing the muscles with a specially designed hammer apparatus, and the frozen muscles were chemically analyzed. Comparable myothermal measurments were made on frogs from the same batch. Results of these experiments indicate: (a) The energy liberation parallels the PC and ATP breakdown with a proportionality constant of 10.7 kcal/mol; (b) comparably designed experiments with sartorius muscles of R. temporaria revealed that the ratio of energy liberation to PC splitting was significantly greater than that observed in R. pipiens sartorius muscles; (c) there is no systematic difference between experiments in which metabolism was arrested by the hammer apparatus and others using a conventional immersion technique.     

Biomechanical behavior of muscle-tendon complex during dynamic human movements

This paper reviews the research findings regarding the force and length changes of the muscle-tendon complex during dynamic human movements, especially those using ultrasonography and computer simulation. The use of ultrasonography demonstrated that the tendinous structures of the muscle-tendon complex are compliant enough to influence the biomechanical behavior (length change, shortening velocity, and so on) of fascicles substantially. It was discussed that the fascicles are a force generator rather than a work generator; the tendinous structures function not only as an energy re-distributor but also as a power amplifier, and the interaction between fascicles and tendinous structures is essential for generating higher joint power outputs during the late pushoff phase in human vertical jumping. This phenomenon could be explained based on the force-length/velocity relationships of each element (contractile and series elastic elements) in the muscle-tendon complex during movements. Through computer simulation using a Hill-type muscle-tendon complex model, the benefit of making a countermovement was examined in relation to the compliance of the muscle-tendon complex and the length ratio between the contractile and series elastic elements. Also, the integral roles of the series elastic element were simulated in a cyclic human heel-raise exercise. It was suggested that the storage and reutilization of elastic energy by the tendinous structures play an important role in enhancing work output and movement efficiency in many sorts of human movements.     

Cyclic nucleotide levels during carbachol-induced smooth muscle contractions.

Cyclic nucleotide levels and tension were measured at various times during carbachol-induced smooth muscle contractions. Cyclic GMP levels were markedly increased during contractions of rat vas deferens, guinea pig myometrium and guinea pig taenia coli, but were unchanged during contractions of rat uterus or guinea pig ileum. No significant changes in cyclic GMP levels could be detected in estrogen-primed rat uteri at any of the times or drug concentrations studied. Even in tissues in which large increases in cyclic GMP levels could be detected during carbachol-induced contractions (i.e. guinea pig myometrium and taenia coli) the contractions appeared to precede the cyclic GMP increases by several seconds. No significant changes in cyclic AMP levels were observed during carbachol-induced contractions in any of the smooth muscles studied. Thus, changes in tissue levels of the cyclic nucleotides do not appear to be responsible for the initiation of carbachol-induced smooth muscle contractions.     

Phosphofructokinase activity and acidosis during short-term tetanic contractions.

Anaerobic energy production is essential for the production of muscular tension when the demand for energy is greater than can be provided aerobically and when oxygen is in short supply. The largest source of anaerobic energy is from the glycolytic pathway. With sustained tetanic contractions, muscle glycolytic activity is high and hydrogen ions (H+) accumulate while tension production decreases. The increasing [H+] and decreasing tension led to the suggestion that H+ inhibits the activity of the regulatory glycolytic enzyme phosphofructokinase (PFK). Early in vitro work confirmed the H+ sensitivity of PFK in the test tube, indicating that little PFK activity should persist at a pH of 6.9-7.0. However, in situ and in vivo experiments suggested that significant PFK activity was maintained during intense contractions when muscle pH decreased to 6.4-6.6. There are several concerns associated with the application of in vitro findings to in vivo exercise situations: (i) there is little in vitro work in mammalian skeletal muscle with substrate and modulator concentrations representative of exercise, (ii) most in vitro analyses of PFK activity are performed following the dilution of the enzyme in mediums with low protein concentration, and (iii) do the modulators identified in vitro exist in high enough in vivo concentrations at rest and during exercise to contribute to the regulation of PFK? More recent in vitro and in situ PFK experiments have overcome some of these concerns. They confirm that during intense, short-term tetanic contractions, PFK activity is well matched to the ATP demand despite decreases in pH to approximately 6.4-6.5.(ABSTRACT TRUNCATED AT 250 WORDS)     

Task failure during fatiguing contractions performed by humans.

By comparing the physiological adjustments that occur when two similar fatiguing contractions are performed to failure, it is possible to identify mechanisms that limit the duration of the more difficult task. This approach has been used to study two fatiguing contractions, referred to as the force and position tasks, which differed in the type of feedback given to the subject and the amount of support provided by the surroundings. Even though the two tasks required a similar net muscle torque during submaximal isometric contractions, the duration that the position task could be sustained was consistently much briefer than that for the force task. The position task involved a greater rate of increase in EMG activity and more marked changes in motor unit recruitment and rate coding compared with the force task. These observations are consistent with the hypothesis that the motor unit pool was recruited more rapidly during the position task. The difference in motor unit behavior appeared to be caused by variation in synaptic input, likely involving heightened sensitivity of the stretch reflex during the position task. Upon repeat performances of the two fatiguing contractions, some subjects were able to increase the time to failure for the force task but not the position task. Furthermore, the time to failure for the position task could be influenced by the postural demands associated with maintaining the position of the limb, and the difference in the two durations was enhanced when the postural activity evoked a pressor response. These observations indicate that the difference in the duration of the two fatiguing contractions was attributable to differences in the control strategy used to sustain the tasks and the magnitude of the associated postural activity.     

Superficial aponeurosis of human gastrocnemius is elongated during contraction: implications for modeling muscle-tendon unit.

Two questions were addressed in this study: (1) how much strain of the superficial aponeurosis of the human medial gastrocnemius muscle (MG) was obtained during voluntary isometric contractions in vivo, (2) whether there existed inhomogeneity of the strain along the superficial aponeurosis. Seven male subjects, whose knees were extended and ankles were flexed at right angle, performed isometric plantar flexion while elongation of superficial aponeurosis of MG was determined from the movements of the intersections made by the superficial aponeurosis and fascicles using ultrasonography. The strain of the superficial aponeurosis at the maximum voluntary contraction, estimated from the elongation and length data, was 5.6+/-1.2%. There was no significant difference in strain between the proximal and distal parts of the superficial aponeurosis. Based on the present result and that of our previous study for the same subjects (J. Appl. Physiol 90 (2001) 1671), a model was formulated for a contracting uni-pennate muscle-tendon unit. This model, which could be applied to isometric contractions at other angles and therefore of wide use, showed that similar strain between superficial and deep aponeuroses of MG contributed to homogeneous fascicle length change within MG during contractions. These findings would contribute to clarifying the functions of the superficial aponeurosis and the effects of the superficial aponeurosis elongation on the whole muscle behavior.     

Expression of the bilateral deficit during reflexively evoked contractions.

During maximal contractions, the sum of forces exerted by homonymous muscles unilaterally is typically larger than the sum of forces exerted by the same muscles bilaterally. This phenomenon is known as the bilateral deficit (BLD), and it is suggested that this deficit is due to neural inhibition. It remains unclear, however, whether such inhibition is mediated by supraspinal mechanisms or by reflex pathways at the level of spinal cord. To further study the origin of likely neural influences, we tested for the presence of BLD under the condition of reflexive force generation. Force output and integrated electromyogram (iEMG) (quadriceps femoris) were measured in 17 male participants after initiation of the myotatic patellar reflex under unilateral and bilateral conditions. A significant BLD of 9.26 +/- 1.19 (P = 0.004) and 16.76 +/- 4.69% (P = 0.001) was found for force and iEMG, respectively. However, because similar findings were not evident during maximal isometric knee extensions, it is difficult to predict the contribution of a spinal mechanism to the BLD under the condition of maximal voluntary activation.     

Effect of calcium ions on rigor contractions in skinned frog muscle fibers

Rigor contractions were examined in skinned frog muscle fibers. The concentrations of calcium ions, pCa = 9.0?5.0, in the solutions which caused rigor were shown to affect the magnitude and time course of the contractions.     

Error propagation from kinematic data to modeled muscle-tendon lengths during walking

Kinematic data from 3D gait analysis together with musculoskeletal modeling techniques allow the derivation of muscle-tendon lengths during walking. However, kinematic data are subject to soft tissue artifacts (STA), referring to skin marker displacements during movement. STA are known to significantly affect the computation of joint kinematics, and would therefore also have an effect on muscle-tendon lengths which are derived from the segmental positions. The present study aimed to introduce an analytical approach to calculate the error propagation from STA to modeled muscle-tendon lengths. Skin marker coordinates were assigned uncorrelated, isotropic error functions with given standard deviations accounting for STA. Two different musculoskeletal models were specified; one with the joints moving freely in all directions, and one with the joints constrained to rotation but no translation. Using reference kinematic data from two healthy boys (mean age 9 y 5 m), the propagation of STA to muscle-tendon lengths was quantified for semimembranosus, gastrocnemius and soleus. The resulting average SD ranged from 6% to 50% of the normalized muscle-tendon lengths during gait depending on the muscle, the STA magnitudes and the musculoskeletal model. These results highlight the potential impact STA has on the biomechanical analysis of modeled muscle-tendon lengths during walking, and suggest the need for caution in the clinical interpretation of muscle-tendon lengths derived from joint kinematics.     

Reliability of surface EMG during sustained contractions of the quadriceps.

The purpose of this study was to determine test-retest reliability for median frequency (MDF) and amplitude of surface EMG during sustained fatiguing contractions of the quadriceps. Twenty-two healthy subjects (11 males and 11 females) were tested on two days held one week apart. Surface EMG was recorded from rectus femoris (RF), vastus lateralis (VL) and vastus medialis (VM) during sustained isometric contractions at 80% and 20% of maximal voluntary contraction (MVC) held to exhaustion. Quadriceps fatigue was described using four measures for both MDF and amplitude of EMG: initial, final, normalized final and slope. For both MDF and amplitude, the initial, final and normalized EMG showed moderate to high reliability for all three muscle groups at both contraction levels (ICC=0.59-0.88 for MDF; ICC=0.58-0.99 for amplitude). Slope of MDF and amplitude was associated with a large degree of variability and low ICCs for the 80% but not the 20% MVC. MDF and amplitude of EMG during sustained contractions of the quadriceps are reproducible; normalized final values of MDF and amplitude show better reliability than slope.