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Baroreflex responsiveness is maintained during isometric exercise in humans.

The simultaneous rise in heart rate and arterial pressure during isometric handgrip exercise suggests that arterial baroreflex control may be altered. We applied incremental intensities of neck suction and pressure to nine healthy young men to alter carotid sinus transmural pressure. Carotid stimuli were delivered during 1) supine control, 2) "anticipation" of beginning exercise, and 3) handgrip (20% of maximum voluntary contraction). Anticipation was a quiet period, immediately preceding the beginning of handgrip, when no muscular work was being performed. Compared with control, the R-R interval prolongation and mean arterial pressure decline provoked by carotid stimuli were decreased during the anticipation period. These data suggest that influences from higher central neural locations may alter baroreflex function. Furthermore, we derived stimulus-response curves relating carotid sinus transmural pressure to changes in R-R interval and mean arterial pressure. These curves were shifted during handgrip; however, calculated regression slopes were not changed from control. The data indicate that isometric handgrip exercise has a specific influence on human carotid baroreflex control of arterial pressure and heart period: baroreflex function curves are shifted rightward during handgrip, whereas baroreflex sensitivity is unchanged. Furthermore, central neural influences may be partially involved in these alterations.

Adult↗

Myoelectrical and mechanical changes linked to length specificity during isometric training.

Mechanical and neural activation changes that accompanied muscle isometric training were studied in males. Training and testing sessions consisted of right elbow isometric flexions. Each experimental group was trained during 5 wk at one of the following angles: 25, 80, and 120 degrees. Bipolar surface electromyogram (EMG) was recorded from the biceps brachii and brachioradialis muscles. An improvement of maximal voluntary contraction (MVC) was always found at the training angle and was systematically greater than at the other angles. Moreover, the shorter the muscle length at which the training has been carried out, the more the gain was limited to the training angle. An increase of the maximal integrated EMG of both biceps brachii and brachioradialis frequently accompanied the improvement of MVC at the training angle.

Adult↗

Simultaneous potentiation and fatigue in quadriceps after a 60-second maximal voluntary isometric contraction.

Potential mechanisms of fatigue (metabolic factors) and potentiation (phosphate incorporation by myosin phosphorylatable light chains) were investigated during recovery from a 60-s maximal voluntary isometric contraction (MVC) in the quadriceps muscle of 12 subjects. On separate days before and for 2 h after the 60-s MVC, either a 1-s MVC or electrically stimulated contractions were used as indexes to test muscle performance. Torque at the end of the 60-s MVC was 57% of the initial level, whereas torques from a 1-s MVC and 50-Hz stimulation were most depressed in the immediate recovery period. At this time, muscle biopsy analyses revealed significant decreases in ATP and phosphocreatine and a 19-fold increase in muscle lactate. Conversely, isometric twitch torque and torque from a 10-Hz stimulus were the least depressed of six contractile indexes and demonstrated potentiation of 25 and 34%, respectively, by 4 min of recovery (P less than 0.05). At this time, muscle lactate concentration was still 16 times greater than at rest. An increased phosphate content of the myosin phosphorylatable light chains (P less than 0.05) was also evident both immediately and 4 min after the 60-s MVC. We conclude that the 60-s MVC produced marked force decreases likely due to metabolic displacement, while the limited decline in the twitch and 10-Hz torques and their significant potentiation suggested that myosin phosphorylation may provide a mechanism to enhance contractile force under conditions of submaximal activation during fatigue.

Adenosine Triphosphate↗

Bilateral deficit is larger for step than for ramp isometric contractions.

The force produced by a muscle group during a maximum effort bilateral task has been reported to be less than that produced during a maximum effort unilateral task. It was hypothesized that if decreased activation of primarily fast motor units is responsible for this bilateral deficit, 1) the bilateral deficit would be larger in step than in ramp isometric contractions and 2) the rate of torque generation in step contractions would be slower in bilateral than in unilateral tasks. Twelve healthy male subjects performed unilateral and bilateral isometric knee joint extension tasks in which maximum torque was generated under ramp and step conditions. The maximum torques produced in the bilateral tasks were 17.0 and 24.6% less than those produced in the unilateral tasks during the ramp and step conditions, respectively (both P < 0.001). The bilateral torque deficit for the step condition was significantly greater than that for the ramp condition (P = 0.012). The rate of torque generation in the step condition was 19.7% slower in the bilateral than in the unilateral task (P = 0.010). Thus the results from these torque measurements support the notion that decreased activation of primarily fast motor units may be responsible for the bilateral deficit.

Adult↗

Role of tendon properties on the dynamic performance of different isometric muscles.

The effect of the tendon's viscoelastic stiffness on the dynamic performance of muscles with different architecture was determined using the cat's medial gastrocnemius and extensor digitorum longus. Dynamic response models were derived under sinusoidal contraction-relaxation in the range of 0.4-6.0 Hz and between 20 and 80% of the muscles' maximal isometric tension, manipulated by orderly recruitment-derecruitment of motor units together with firing rate increase-decrease. It was shown that, for isometric contractions at the muscle's optimum length, the dynamic response of the muscles was not significantly different before and after dissection of the tendon. Therefore the conclusion that under these conditions the tendon acts like a stiff force transmitter without significantly modifying the muscle's performance was confirmed and extended to muscles with different architecture.

Action Potentials↗

Decreased Ca2+ sensitivity of isometric tension in skinned cardiac myocytes from tail-suspended rats.

Tail suspension in rats causes a cephalic shift in blood, resulting in a volume load on the heart similar to that observed during microgravity spaceflight or mild heart failure. The present study determined the influence of increased cardiac hemodynamic load on myofilament isometric tension as a function of Ca2+ concentration in skinned cardiac myocytes of control and 7-day head-down tilt Sprague-Dawley rats. Isometric force of single skinned myocytes was measured by attaching cells with adhesive to a force transducer and piezoelectric translator. A significant decrease in the Ca2+ sensitivity of tension was observed in cardiac myocytes from suspended rats [pCa of half-maximal tension (pCa50) of 5.83 +/- 0.03] compared with control rats (pCa50 of 5.94 +/- 0.03). Maximum tension generation and slope of the tension-pCa relationship were unaffected by head-down tilt. Electrophoretic analysis of myofilament proteins indicates differences in expression of proteins in the 50-60 and 100-120 kDa ranges; immunoblot analysis of tubulin (50 kDa) expression indicates no change in the ratio of beta-tubulin to light chain 1 or tropomyosin. Decreased force-producing ability at a given submaximum Ca2+ concentration in cardiac myocytes from suspended rats suggests a decrease in contractility possibly due to changes in cardiac myofilament protein expression following chronic elevated volume load on the heart.

Animals↗

Mechanical and metabolic determination of VO2 and fatigue during repetitive isometric contractions in situ.

Repetitive isometric tetanic contractions (1/s) of the canine gastrocnemius-plantaris muscle were studied either at optimal length (Lo) or short length (Ls; approximately 0.9 . Lo), to determine the effects of initial length on mechanical and metabolic performance in situ. Respective averages of mechanical and metabolic variables were (Lo vs. Ls, all P < 0.05) passive tension (preload) = 55 vs. 6 g/g, maximal active tetanic tension (Po) = 544 vs. 174 (0.38 . Po) g/g, maximal blood flow (Q) = 2.0 vs. 1.4 ml . min-1 . g-1, and maximal oxygen uptake (VO2) = 12 vs. 9 micromol . min-1 . g-1. Tension at Lo decreased to 0.64 . Po over 20 min of repetitive contractions, demonstrating fatigue; there were no significant changes in tension at Ls. In separate muscles contracting at Lo, Q was set to that measured at Ls (1.1 ml . min-1 . g-1), resulting in decreased VO2 (7 micromol . min-1 . g-1), and rapid fatigue, to 0.44 . Po. These data demonstrate that 1) muscles at Lo have higher Q and VO2 values than those at Ls; 2) fatigue occurs at Lo with high VO2, adjusting metabolic demand (tension output) to match supply; and 3) the lack of fatigue at Ls with lower tension, Q, and VO2 suggests adequate matching of metabolic demand, set low by short muscle length, with supply optimized by low preload. These differences in tension and VO2 between Lo and Ls groups indicate that muscles contracting isometrically at initial lengths shorter than Lo are working under submaximal conditions.

Animals↗

Nonisometric behavior of fascicles during isometric contractions of a human muscle.

Fascicle length, pennation angle, and tendon elongation of the human tibialis anterior were measured in vivo by ultrasonography. Subjects (n = 9) were requested to develop isometric dorsiflexion torque gradually up to maximal at the ankle joint angle of 20 degrees plantarflexion from the anatomic position. Fascicle length shortened from 90 +/- 7 to 76 +/- 7 (SE) mm, pennation angle increased from 10 +/- 1 to 12 +/- 1 degrees, and tendon elongation increased up to 15 +/- 2 mm with graded force development up to maximum. The tendon stiffness increased with increasing tendon force from 10 N/mm at 0-20 N to 32 N/mm at 240-260 N. Young's modulus increased from 157 MPa at 0-20 N to 530 MPa at 240-260 N. It can be concluded that, in isometric contractions of a human muscle, mechanical work, some of which is absorbed by the tendinous tissue, is generated by the shortening of muscle fibers and that ultrasonography can be used to determine the stiffness and Young's modulus for human tendons.

Ankle Joint↗

Effect of muscle mass and intensity of isometric contraction on heart rate.

The purpose of this study was to determine the effect of muscle mass and the level of force on the contraction-induced rise in heart rate. We conducted an experimental study in a sample of 28 healthy men between 20 and 30 yr of age (power: 95%, alpha: 5%). Smokers, obese subjects, and those who performed regular physical activity over a certain amount of energetic expenditure were excluded from the study. The participants exerted two types of isometric contractions: handgrip and turning a 40-cm-diameter wheel. Both were sustained to exhaustion at 20 and 50% of maximal force. Twenty-five subjects finished the experiment. Heart rate increased a mean of 15.1 beats/min [95% confidence interval (CI): 5.5-24.6] from 20 to 50% handgrip contractions, and 20.7 beats/min (95% CI: 11.9-29.5) from 20 to 50% wheel-turn contractions. Heart rate also increased a mean of 13.3 beats/min (95% CI: 10.4-16.1) from handgrip to wheel-turn contractions at 20% maximal force, and 18.9 beats/min (95% CI: 9. 8-28.0) from handgrip to wheel-turn contractions at 50% maximal force. We conclude that the magnitude of the heart rate increase during isometric exercise is related to the intensity of the contraction and the mass of the contracted muscle.

Adult↗

Blunted pressor and intramuscular metabolic responses to voluntary isometric exercise in multiple sclerosis.

To test the hypothesis that a lower mean arterial pressure (MAP) response during voluntary isometric exercise in multiple sclerosis (MS) is related to a dampened muscle metabolic signal, 9 MS and 11 control subjects performed an isometric dorsiflexor contraction at 30% maximal voluntary contraction until target failure (endurance time). We made continuous and noninvasive measurements of heart rate and MAP (Finapres) and of intramuscular pH and P(i) (phosphorus magnetic resonance spectroscopy) in a subset of 6 MS and 10 control subjects. Endurance times and change in heart rate were similar in MS and control subjects. The decrease in pH and increase in P(i) were less throughout exercise in MS compared with control subjects, as was the change in MAP response. Differences in muscle strength accounted for some of the difference in MAP response between groups. Cardiovascular responses during Valsalva and cold pressor tests were similar in MS and control subjects, suggesting that the blunted MAP response during exercise in MS was not due to a generalized dysautonomia. The dampened metabolic response in MS subjects was not explained by inadequate central muscle activation. These data suggest that the blunted pressor response to exercise in MS subjects may be largely appropriate to a blunted muscle metabolic response and differences in contracting muscle mass.

Adult↗

Terminology for contractions of muscles during shortening, while isometric, and during lengthening.

Communication among scientists must be clear and concise to avoid ambiguity and misinterpretations. The selection of words must be based on accepted definitions. The fields of biomechanics, muscle physiology, and exercise science have had a particularly difficult time with terminology, arising from the complexity of muscle contractions and by the use of inappropriate terminology by scientists. The dictionary definition of the verb "contract," specifically for the case of muscle, is "to undergo an increase in tension, or force, and become shorter." Under all circumstances, an activated muscle generates force, but an activated muscle generating force does not invariably shorten! During the 1920s and 1930s, investigators recognized that the interaction between the force generated by the muscle and the load on the muscle results in either shortening, no length change (isometric), or lengthening of the muscle. The recognition that muscles perform three different types of "contractions" required that contraction be redefined as "to undergo activation and generate force." Modifiers of contraction are then needed to clarify the lack of movement or the directionality of movement. Despite the contradiction, for 75 years the lack of movement has been termed an "isometric contraction." The directionality of the movement is then best described by the adjectives "shortening" and "lengthening." The definitions of "concentric" as "having the same center" and of "eccentric" as "not having the same center" are consistent with hypertrophy, or remodeling of the heart muscle, but are inappropriate to describe the contractions of skeletal muscles.

Animals↗

Effect of quadriceps femoris muscle length on neural activation during isometric and concentric contractions.

The effect of muscle length on neural drive (here termed "neural activation") was investigated from electromyographic activities and activation levels (twitch interpolation). The neural activation was measured in nine men during isometric and concentric (30 and 120 degrees /s) knee extensions for three muscle lengths (35, 55, and 75 degrees knee flexion, i.e., shortened, intermediate, and lengthened muscles, respectively). Long (76 degrees ), medium (56 degrees ), and short (36 degrees ) ranges of motion were used to investigate the effect of the duration of concentric contraction. Neural activation was found to depend on muscle length. Reducing the duration of contraction had no effect. Neural activation was higher with short muscle length during isometric contractions and was weaker for shortened than for intermediate and lengthened muscles performing 120 degrees /s concentric contractions. Muscle length had no effect on 30 degrees /s concentric neural activation. Peripheral mechanisms and discharge properties of the motoneurons could partly explain the observed differences in the muscle length effect. We thus conclude that muscle length has a predominant effect on neural activation that would modulate the angular velocity dependency.

Adult↗

Neuromuscular fatigue development during maximal concentric and isometric knee extensions.

This study aimed to investigate mechanisms of neuromuscular fatigue during maximal concentric and isometric leg extensions inducing similar torque decrements. Nine physically active men performed two separate fatiguing sessions maintained until similar torque decreases were obtained. The first session, only conducted under isokinetic concentric conditions (CON), consisted of three series of 30 maximal voluntary concentric knee extensions (60 degrees/s). The second session, exclusively isometric (ISO), mimicked the torque decreases registered during the CON session while performing three long-lasting ISO contractions. Maximal voluntary torque, activation level (twitch interpolation technique), electromyographic activity (root mean square and median frequency) of the vastus lateralis muscle, and electrically evoked doublet-twitch mechanical properties were measured before and at the end of each of the three series. After the three series, similar torque decrements were obtained for both fatiguing procedures. The total fatiguing contraction durations were not different among procedures. With equivalent voluntary torque decrements, the doublet-twitch amplitude reduction was significantly greater (P<0.01) during the two first series of the CON procedure compared with ISO. No difference was observed for the third series. Although no difference was recorded with fatigue for median frequency changes between CON and ISO, activation levels and root mean square values demonstrated greater reductions (P<0.05) for all three series during the ISO procedure compared with CON. Performing CON or ISO fatiguing exercises demonstrated different fatigue origins. With CON exercises, peripheral fatigue developed first, followed by central fatigue, whereas with ISO exercises the fatigue pattern was inverted.

Adult↗

Postactivation potentiation in a human muscle: effect on the rate of torque development of tetanic and voluntary isometric contractions.

Postactivation potentiation (PAP), a mechanism by which the torque of a muscle twitch is increased following a conditioning contraction, is well documented in muscular physiology, but little is known about its effect on the maximal rate of torque development and functional significance during voluntary movements. The objective of this study was to investigate the PAP effect on the rate of isometric torque development of electrically induced and voluntary contractions. To that purpose, the electromechanical responses of the thumb adductor muscles to a single electrical stimulus (twitch), a train of 15 pulses at 250 Hz (HFT(250)), and during ballistic (i.e., rapid torque development) voluntary contractions at torque levels ranging from 10 to 75% of maximal voluntary contraction (MVC) were recorded before and after a conditioning 6-s MVC. The results showed that the rate of torque development was significantly (P < 0.001) increased after the conditioning MVC, but the effect was greater for the twitch ( approximately 200%) compared with the HFT(250) ( approximately 17%) or ballistic contractions (range: 9-24%). Although twitch potentiation was maximal immediately after the conditioning MVC, maximal potentiation for HFT(250) and ballistic contractions was delayed to 1 min after the 6-s MVC. Furthermore, the similar degree of potentiation for the rate of isometric torque development between tetanic and voluntary ballistic contractions indicates that PAP is not related to the modality of muscle activation. These observations suggest that PAP may be considered as a mechanism that can influence our contractions during daily tasks and can be utilized to improve muscle performance in explosive sports.

Adaptation, Physiological↗

Firing rates of motor units in human vastus lateralis muscle during fatiguing isometric contractions.

We investigated the firing rate of motor units in the vastus lateralis muscle in five healthy young men (mean = 21.4 yr, SD = 0.9) during a sequence of isometric constant-torque contractions repeated to exhaustion. The contractions were sustained at 20% of the maximal voluntary level, measured at the beginning of the test sequence. Electromyographic (EMG) signals were recorded via quadrifilar fine-wire electrodes and subsequently decomposed into their constituent motor unit action potentials to obtain the motor unit firing times. In addition, we measured the whole muscle mechanical properties during the fatigue task using electrical stimulation. The firing rate of motor units first decreased within the first 10-20% of the endurance time of the contractions and then increased. The firing rate increase was accompanied by recruitment of additional motor units as the force output remained constant. The elicited twitch and tetanic torque responses first increased and then decreased. The two processes modulated in a complementary fashion at the same time. Our data suggest that, when the vastus lateralis muscle is activated to maintain a constant torque output, its motoneuron pool receives a net excitatory drive that first decreases to compensate for the short-lived potentiation of the muscle force twitch and then increases to compensate for the diminution of the force twitch. The underlying inverse relationship between the firing rate and the recruitment threshold that has been reported for nonfatigued contractions is maintained. We, therefore, conclude that the central nervous system control of vastus lateralis motor units remains invariant during fatigue in submaximal isometric isotonic contractions.

Action Potentials↗

Muscle activity determined by cosine tuning with a nontrivial preferred direction during isometric force exertion by lower limb.

We investigated how the CNS selects a unique muscle activation pattern under a redundant situation resulting from the existence of bi-articular muscles. Surface electromyographic (EMG) activity was recorded from eight lower limb muscles while 11 subjects were exerting isometric knee and hip joint torque simultaneously (T(k) and T(h), respectively. Extension torque was defined as positive). The knee joint was kept at either 90 or 60 degrees. Various combinations of torque were imposed on both joints by pulling a cable attached to an ankle brace with approximately three levels of isometric force in 16 directions. The distribution of the data in the three-dimensional plot (muscle activation level quantified by the root mean squared value of EMG vs. T(k) and T(h)) demonstrates that the muscle activation level M can be approximated by a single model as M = left flooraT(k) + bT(h) right floor where left floorx right floor = max (x,0) and a and b are constants. The percentage of variance explained by this model averaged over all muscles was 82.3 +/- 14.0% (mean +/- SD), indicating that the degree of fit of the data to the plane was high. This model suggests that the CNS uses a cosine tuning function on the torque plane (T(k), T(h)) to recruit muscles. Interestingly, the muscle's preferred direction (PD) defined as the direction where it is maximally active on the torque plane deviated from its own mechanical pulling direction (MD). This deviation was apparent in the mono-articular knee extensor (MD = 0 degrees , whereas PD = 14.1 +/- 3.7 degrees for vastus lateralis) and in the mono-articular hip extensor (MD = 90 degrees, whereas PD = 53.4 +/- 6.4 degrees for gluteus maximus). Such misalignment between MD and PD indicates that the mono-articular muscle's activation level depends on the torque of the joint that it does not span. Practical implications of this observation for the motor control studies were discussed. We also demonstrated that the observed shift from the MD to the PD is plausible in the configuration of our musculo-skeletal system and that the experimental results are likely to be explained by the CNS process to minimize the variability of the endpoint force vector under the existence of signal-dependent noise.

Adult↗

Rotation of motoneurons during prolonged isometric contractions in humans.

Prolonged and weak isometric contractions can result in neuromuscular fatigue. Alternation of discharge of motor units with similar thresholds (termed rotation) could be useful to minimize neuromuscular fatigue by providing periods for metabolic recovery of the contractile elements. In the present study, we investigated the prevalence of motoneuron rotation during prolonged contractions of distal limb muscles. Electromyographic (EMG; needle and surface) was recorded from muscles of the forearm and distal leg. The subject made a slowly increasing isometric contraction to recruit and discharge a motor unit (1) for a prolonged period of time (> 30 min). Sometimes an additional motor unit (2) was recruited in which case the subject relaxed the contraction slightly so that only one motor unit remained tonic. Often it was this newly recruited motor unit (i.e., unit 2) that continued discharging, while motor unit 1 fell silent. Continued contraction would then lead to the resumption of tonic discharge of unit 1 and silence of unit 2. This would complete a rotation between motor units 1 and 2. During prolonged contractions, rotation was observed in approximately 80% of the motor-unit pairs examined. There was no difference in rotation incidence by muscle type. For the unit pairs showing rotation, surface EMG values were significantly higher immediately prior to rotation than after rotation had occurred. Our findings show that rotation of motor units with similar recruitment thresholds during such contractions is common in distal muscles of the arm and leg and may help offset neuromuscular fatigue.

Adult↗

Discharge properties of primate forearm motor units during isometric muscle activity.

Activity of single motor units (MUs) was recorded in forelimb muscles of rhesus macaques while they generated isometric ramp-and-hold torques about the wrist. Multiunit electromyographic (EMG) activity was recorded from 10-12 identified flexor and extensor muscles of the wrist and digits with implanted EMG wire electrodes. Single MUs from these muscles were recorded with a remotely controlled tripolar microelectrode array. The parent muscle of each MU was determined by compiling MU-triggered averages of multiunit EMGs. The MU firing patterns during the isometric task were determined from response histograms aligned with change in torque. At moderate torque levels, MUs (n = 86) exhibited four types of discharge patterns during the ramp-and-hold trajectory: phasic-tonic (23%), tonic (33%), decrementing (39%), and phasic (5%). Phasic-tonic MUs exhibited a phasic burst of activity during the torque ramp which exceeded the firing rate during the static hold period. Both phasic-tonic and tonic MUs exhibited a constant mean firing rate during the hold period; the discharge of decrementing MUs gradually decreased during the static hold. Phasic MUs fired only during the change in force. The relation between MU firing rate and torque was investigated as the monkeys generated responses of different levels of static torque during the hold period. Mean firing rate during the hold was found to be proportional to static torque up to moderate torque levels, where it reached a maximum. In the linear range, the mean rate-torque slope was 3.4 +/- 1.9 imp/s per 10(5) dyn . cm (n = 9).

Animals↗