Mechanical properties of aponeurosis and tendon of the cat soleus muscle during whole-muscle isometric contractions

Recent studies have suggested that the mechanical properties of aponeurosis are not similar to the properties of external tendon. In the present study, the lengths of aponeurosis, tendon, and muscle fascicles were recorded individually, using piezoelectric crystals attached to the surface of each structure during isometric contractions in the cat soleus muscle. We used a surgical microscope to observe the surface of the aponeurosis, which revealed a confounding effect on measures of aponeurosis length due to sliding of a thin layer of epimysium over the proximal aponeurosis. After correcting for this artifact, the stiffness computed for aponeurosis was similar to tendon, with both increasing from around 8 F₀/L_c (F₀ is maximum isometric force and L_c is tissue length) at 0.1 F₀ to 30 F₀/L_c at forces greater than 0.4 F₀. At low force levels only (0.1 F₀), aponeurotic stiffness increased somewhat as fascicle length increased. There was a gradient in the thickness of the aponeurosis along its length: its thickness was minimal at the proximal end and maximal at the distal end, where it converged to form the external tendon. This gradient in thickness appeared to match the gradient in tension transmitted along this structure. We conclude that the specific mechanical properties of aponeurosis are similar to those of tendon. © 1995 Wiley-Liss, Inc.     

The effect of prolonged isometric contractions on muscle fluid balance

Ultrasound scanning was performed at three sites above the fossa supraspinata on nine healthy subjects and five patients with myofascial shoulder pain. This method produced a well-defined depiction of the soft tissue layers above the fossa supraspinata and reproducible muscle thickness measurements. In the healthy subjects the average distance from the skin surface to the trapezius muscle was 7.7 mm and the average thickness of the trapezius muscle was 5.3 mm, and the average thickness of supraspinatus muscle was 20.0 mm. The supraspinatus muscle was thinner at the medial measuring site than at the other two sites. In contrast, a tendency towards a larger distance was seen from the skin to trapezius muscle at the medial measuring site than at the other two sites. No statistical differences were found between the two groups of subjects either at rest or during brief shoulder abductions. All the subjects performed a 30° unilateral isometric shoulder abduction test to exhaustion. The median endurance time was 33 min for the healthy subjects and only 5 min for the patients. The ratings of perceived exertion (RPE) were in line with this, since the increment in RPE with time was larger for the patients than for the healthy group. The reduced shoulder abduction endurance time in the patient group may have been related to impaired muscle function and/or pain development. During the 33-min shoulder abduction in the healthy subjects, the thickness of supraspinatus muscle increased by 14%, indicating muscle swelling, whereas the thickness of trapezius muscle remained constant. The fluid imbalance in the supraspinatus muscle compartment may well play a role in the development of muscle fatigue and the disorders found in industry resulting from prolonged work with arms elevated.     

Sarcomere dynamics in skeletal muscle myofibrils during isometric contractions

The main goal of this study was to evaluate the dynamics of sarcomeres during isometric activation of skeletal muscle myofibrils. Rabbit psoas myofibrils (n=14) were attached between a pair of cantilevers for force measurements at one side and a rigid glass needle at the other side, and their images were used for measurements of individual sarcomere lengths (SL) during contractions. Myofibrils were set at average SL between 2.13 and 3.06 μm, and were activated and held isometric for 20–35 s during which SL and force were continuously measured. SL dispersion increased from the rest state to activation, but it remained mostly constant during the activation period. Even with the length non-uniformity developed during myofibril activation, most sarcomeres stabilized their length changes during the isometric contraction. As a result, sarcomeres contracted at different degrees of filament overlap while producing similar forces. When the myofibrils were separated in two groups that produced force at averaged short (≤2.5 μm) or long (≥2.5 μm) SL, the initial non-uniformity was greater in long lengths, but changes observed in sarcomeres during the activation period were similar, suggesting that sarcomere stability is not length-dependent.     

Functional subdivision of the upper trapezius muscle during maximal isometric contractions

The electromyographic (EMG) activity pattern across the upper trapezius of 22 healthy subjects was investigated during maximal isometric contractions. Eight bipolar surface electrodes with 10 mm distance between adjacent electrode pairs were placed on a line from the clavicle to the scapula. At the region near the clavicle the highest EMG amplitudes were recorded during 90 ° arm abduction. At the more posterior parts the highest amplitudes were found both during arm abduction and shoulder elevation. A double differential recording technique which reduced the EMG cross-talk contribution supported the finding that the upper trapezius was differently activated when the arm posture was changed. The normalized EMG amplitude-force relationship during the shoulder elevation showed a curvilinear relationship on the anterior part of the upper trapezius with a slower increase in EMG amplitude than force at low force. The slope of the curve, at low force, increased gradually in the posterior direction on the upper trapezius. The EMG activity patterns across the upper trapezius indicate a flexibility in motor activation which maybe reflects a functional optimization of the contractions performed by this muscle.     

Fatigue of mouse diaphragm muscle in isometric and isotonic contractions

Fatiguabilities of mouse diaphragm muscle in vitro in isometric and isotonic contractions were compared in this study. Isolated mouse diaphragm muscle was stimulated repetitively to induce fatigue during both isometric and isotonic contractions. The supramaximal electrical stimulation used was a train of 100-Hz, 0.5-ms pulses delivered to the muscle every 2 s for 0.5 s. The percentage decrease in isometric tension from beginning to end of the fatiguing process was used as the index of fatigue. The experiments were carried out at different PO2 levels in both normal and zero-glucose Ringer solutions. It was found that fatigue developed more rapidly in isotonic contractions than in isometric ones. Also, the extracellular glucose level demonstrated little effect on the muscle's short-term fatiguability, whereas reductions in the extracellular PO2 exerted a profound effect, especially in the case of isotonic fatigue.     

Electromechanical delay in the vastus lateralis muscle during dynamic isometric contractions

Electromechanical delay (EMD) values were obtained using a cross-correlation technique for a series of 14 repetitive submaximal dynamic isometric contractions of the vastus lateralis performed by five subjects. To avoid a phase lag, which is introduced with one-way filtering, the EMG was processed with a bi-directional application of a second-order Butterworth filter. A mean EMD value of 86 ms (SD = 5.1 ms) was found. Moreover, contraction and relaxation delays were computed and compared. There was a significant difference between the contraction and relaxation delays (P less than 0.005). The mean contraction delay was 81.9 ms and the mean relaxation delay was 88.8 ms. Despite this significant difference, the computed contraction and relaxation delay values lie in the same range as the total phase lag, calculated with the cross-correlation technique. The magnitude of EMD values found supports the need to account for this delay when interpreting temporal aspects of patterns of intermuscular coordination.     

Effects of aging on EMG variables during fatiguing isometric contractions

The purpose of this study was to evaluate the neuromuscular adaptation that occurred with aging, by comparing young and aged subjects with respect to changes in surface EMG from the tibialis anterior muscle during fatiguing contractions. EMG variables such as the averaged rectified value (ARV), median frequency (MDF), and muscle fiber conduction velocity (MFCV) were calculated during maximal (MVC, 3 sec) and submaximal (60% MVC, 60 sec) isometric contractions. Muscular force, ARV, MDF, and MFCV during MVC were significantly greater in the young than in the elderly (p < 0.05). EMG amplitude increased and the waveform slowed in all subjects during submaximal contractions, indicating the development of local muscle fatigue. As fatigue progressed, the ARV increased and the MDF and MFCV decreased significantly (p < 0.01). The fatigue-induced changes in the MDF and MFCV were significantly smaller in aged than in young subjects (p < 0.05), a trend also seen in the ARV change, which means that the elderly cannot be fatigued as much as the young with contractions of the same relative intensity. These results as a whole suggest that the aged subjects hold an adaptive motor strategy to cope with age-related neuromuscular deteriorations, due to the decline of motor unit activation and selective atrophy of fast twitch muscle fibers.     

Effect of muscle length on surface EMG wave forms in isometric contractions

To elucidate the influence of muscle length on surface EMG wave form, comparisons were made of surface EMGs of the biceps and triceps brachii muscles during isometric contractions at different muscle lengths. Muscle lengths were altered by setting the elbow joint angle at several intervals between the limits of extension and flexion. The intensity of the isometric contractions was 25% of maximum voluntary contraction at the individual joint angles. Slowing was obvious in the EMG wave forms of biceps as muscle length increased. The so-called 'Piper rhythm' appeared when the muscle was more than moderately lengthened. The slowing trend with muscle lengthening, though less marked, was also seen in triceps. Zero-cross analysis revealed quasi-linear relationships between muscle length and slowing. Frequency analysis confirmed the development of 'Piper rhythm'. An attempt was made to interpret the slowing associated with muscle lengthening in terms of the propagation of myoelectric signals in muscle fibers. given the effect of muscle length on EMG wave forms, a careful control of joint angle may be required in assessing local making fatigue when using EMG spectral indices.     

The effect of muscle fatigue on the isometric contractile characteristics of skeletal muscle in the cat

The rise time of an isometric twitch, the tetanic tension, the twitch tetanus ratio, the frequency-tension relationship, and the height of the MUAP (motor unit action potential) were measured in fast twitch (medial gastrocnemius) and slow twitch (soleus) muscles of the cat immediately before, in the middle, and immediately after fatiguing isometric contractions at tensions of 30, 50 and 80% of each muscle's initial strength (tetanic tension recorded from the unfatigued muscle). Although the twitch-tetanus ratio was always less for the soleus than for the medial gastrocnemius muscles, the twitch-tetanus ratio for any one muscle was constant throughout the duration of fatiguing isometric contractions at any of the tensions examined. In contrast, the twitch tension and tetanic tension of the muscles were both less after the contractions, the largest reduction occurring for both muscles during contractions sustained at the lowest isometric tensions. The time to peak tension of an isometric twitch was prolonged for both muscles following the contractions. This was associated with a corresponding shift in the frequency tension relationship such that at the point of muscular fatigue, the muscles tetanized at lower frequencies of stimulation than did the unfatigued muscle. In contrast, the amplitude of the MUAP showed only a modest reduction throughout the duration of the fatiguing contractions.     

Effects of elastic loads on the contractions of cat muscles

The nerves to plantaris and soleus muscles in the cat were stimulated with maximal single shocks and with random stimulus trains which produced partially fused contractions. In order to obtain information on the mechanism of muscular contraction, the effects of allowing the muscles to shorten against various elastic loads were studied in the time domain and in the frequency domain. When springs of increasing stiffness were placed in series with the muscle, the twitch tension increased greatly. The gain of the frequency response curve was also much greater with stiffer springs. The shape of the frequency response curve for plantaris muscle could usually be described by that expected for a second-order system with two real time constants or rate constants. The rate constants changed in qualitatively similar ways in response to increased stiffness of an elastic load, increased muscle length and increased mean rate of nerve stimulation. These results are in agreement with the hypothesis that the linear responses of muscles working against elastic loads are determined by the values of two rate constants. Thus, of the many processes associated with contraction, only two are rate-limiting: one associated with the viscoelastic properties of muscle and the second associated with the reuptake of Ca into the sarcoplasmic reticulum. Non-linear aspects of muscular contraction are also discussed. These are more prominent in soleus muscle than in plantaris muscle.Graduate student of the Medical Research Council of Canada.Formerly a Post-doctoral Fellow of the Muscular Dystrophy Association of Canada.     

Nonisometric behavior of fascicles during isometric contractions of a human muscle

Fascicle length, pennation angle, and tendonelongation of the human tibialis anterior were measured in vivo byultrasonography. Subjects (n = 9) wererequested to develop isometric dorsiflexion torque gradually up tomaximal at the ankle joint angle of 20° plantarflexion from theanatomic position. Fascicle length shortened from 90 ± 7 to 76 ± 7 (SE) mm, pennation angle increased from 10 ± 1 to 12 ± 1°, and tendon elongation increased up to 15 ± 2 mm with gradedforce development up to maximum. The tendon stiffness increased withincreasing tendon force from 10 N/mm at 0-20 N to 32 N/mm at240-260 N. Young's modulus increased from 157 MPa at 0-20 Nto 530 MPa at 240-260 N. It can be concluded that, in isometriccontractions of a human muscle, mechanical work, some of which isabsorbed by the tendinous tissue, is generated by the shortening ofmuscle fibers and that ultrasonography can be used to determine thestiffness and Young's modulus for human tendons.

     

A comparison of models explaining muscle activation patterns for isometric contractions

One of the main problems in motor-control research is the muscle load sharing problem, which originates from the fact that the number of muscles spanning a joint exceeds the number of degrees of freedom of the joint. As a consequence, many different possibilities exist for the activation of muscles in order to produce a desired joint torque. Several models describing muscle activation have been hypothesized over the last few decades to solve this problem. This study presents theoretical analyses of the various models and compares the predictions of these models with new data on muscle activation patterns for isometric contractions in various directions. None of the existing models fitted the experimental data in all aspects. The best fit was obtained by models based on minimization of the squared sum of muscle forces (∑ _m φ² _m , which is almost equivalent to the Moore-Penrose pseudo-inverse solution), muscle stress σ (∑ _m σ _m ²) or muscle activation α (∑ _m α _m ²). Since muscle activation patterns are different for isometric contractions and for movements, it could well be that other models or optimization criteria are better suited to describe muscle activation patterns for movements. The results of our simulations demonstrate that the predicted muscle activation patterns do not depend critically on the parameters in the model. This may explain why muscle activation patterns are highly stereotyped for all subjects irrespective of differences between subjects in many neuro-anatomical aspects, such as, for example, in the physiological cross-sectional area of muscle. Received: 24 September 1998 / Accepted in revised form: 1 March 1999     

Mechanical behavior of skeletal muscle during intermittent voluntary isometric contractions in humans

Vøllestad, N. K., I. Sejersted, and E. Saugen. Mechanical behavior of skeletal muscle duringintermittent voluntary isometric contractions in humans.J. Appl. Physiol. 83(5):1557-1565, 1997.Changes in contractile speed and force-fusionproperties were examined during repetitive isometric contractions withthe knee extensors at three different target force levels. Sevenhealthy subjects were studied at target force levels of 30, 45, and60% of their maximal voluntary contraction (MVC) force. Repeated 6-s contractions followed by 4-s rest were continued until exhaustion. Contractile speed was determined for contractions elicited by electrical stimulation at 1-50 Hz given during exercise and a subsequent 27-min recovery period. Contraction time remained unchanged during exercise and recovery, except for an initial rapid shift in thetwitch properties. Half relaxation time(RT_1/2) decreased gradually by 20-40% during exercise at 30 and 45% of MVC. In the recovery period, RT_1/2 values werenot fully restored to preexercise levels. During exercise at 60% MVC,the RT_1/2 decreased for twitches and increased for the 50-Hz stimulation. In the recovery period after60% MVC, RT_1/2 values declinedtoward those seen after the 30 and 45% MVC exercise. The forceoscillation amplitude in unfused tetani relative to the mean forceincreased during exercise at 30 and 45% MVC but remained unalteredduring the 60% MVC exercise. This altered force-fusion was closelyassociated with the changes inRT_1/2. The faster relaxation mayat least partly explain the increased energy cost of contractionreported previously for the same type of exercise.

     

Experimental muscle pain increases mechanomyographic signal activity during sub-maximal isometric contractions.

This study was designed to investigate the local effect of experimental muscle pain on the MMG and the surface EMG during a range of sub-maximal isometric contractions. Muscle pain was induced by injections of hypertonic saline into the biceps brachii muscle in 12 subjects. Injections of isotonic saline served as a control. Pain intensity and location, MMG and surface EMG from the biceps brachii were assessed during static isometric (0%, 10%, 30%, 50% and, 70% of the maximal voluntary contraction) and ramp isometric (0-50% of the maximal voluntary contraction) elbow flexions. MMG and surface EMG signals were analyzed in the time and frequency domain. Experimentally induced muscle pain induced an increase in root mean square values of the MMG signal while no changes were observed in the surface EMG. Most likely this increase reflects changes in the mechanical contractile properties of the muscle and indicates compensatory mechanisms, i.e. decreased firing rate and increased twitch force to maintain a constant force output in presence of experimental muscle pain. Under well-controlled conditions, MMG recordings may be more sensitive than surface EMG recordings and clinically useful for detecting non-invasively increased muscle mechanical contributions during muscle pain conditions.