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Changes in reflex excitability following isometric contraction in humans.

Enhancement of muscle stretch following isometric contraction has been thought to occur as a result of inhibitory reflex mechanisms. Experiments with electrical stimulation (H-reflex) have demonstrated maximal H-reflex suppression during force relaxation followed by gradual recovery over the following 20 s. There has been considerable speculation as to whether electrical and mechanical stimulation elicit similar response behaviour. The present study examined postisometric reflex modulation following both stimulation modalities. In ten subjects dorsiflexion stimuli varying in speed and amplitude were applied after 30% and 60% maximal voluntary contraction (MVC). Modulation of the mechanically and electrically evoked responses following isometric plantarflexion was investigated. Reflex responses following both stimulation modalities were depressed during the course of force relaxation. A rather fast recovery was observed in mechanical stimulation. Postisometric response modulation was neither altered by the amount of isometric plantarflexion, nor by the amplitude of the applied stretch stimulus. With increasing velocity of the applied dorsiflexion, however, the shape of the reflex modulation persisted, but the magnitude of the responses was significantly enhanced. In electrical stimulation, however, recovery was delayed. It is suggested that postisometric reflex modulation is due to presynaptic inhibition. Moreover, possible peripheral mechanisms resulting from alpha-gamma-coactivation may also affect the stretch receptor itself because of inherent stiffness properties. The latter possibility particularly would explain the differences between mechanical and electrical stimulus modalities. With respect to practical implications, the very fast recovery (< 400 ms) of the stretch responses to control values strongly contradicts the interpretation that after isometric precontraction, suppression of reflex activity might be used for more efficient stretching of the tendomuscle system.

Adult

Physiological responses to sustained isometric contractions during one- and two-armed work. Role of muscle mass on cardiovascular responses to isometric work.

Cardiovascular and ventilatory responses were investigated during one- and two-armed isometric contractions at the same level of maximum voluntary contraction force (MVC). Significant differences were observed in heart rate, blood pressure, pulse arrival time, and oxygen intake in the contractions as compared with each other. These findings contradict the generally accepted view that cardiovascular responses at the same relative force in isometric muscle contractions, irrespective of the size of muscle mass engaged, are equal.

Adult

Frequency analysis of the surface electromyogram during sustained isometric contractions.

Four male and four female volunteers served as subjects in these experiments to assess the frequency components of the surface EMG during and following brief (3 s) and sustained isometric contractions of the handgrip muscles. Two types of fatiguing contractions were performed. Contractions were either maintained to fatigue at a constant tension of up to 100% of their strength or were maintained as a sustained maximal effort in the unfatigues or previously fatigued muscle. The frequency components of the surface EMG were assessed by calculating the power spectra of 1.5 s samples of the EMG from a fundamental frequency of 4 Hz through the first 128 harmonics by Fourier analysis; the centre frequencies of the resultant power spectra were then used as an index of the mean frequency of the EMG. The results of these experiments showed that the centre frequency was independent of the tension exerted by the muscle during brief isometric contractions but decreased linearly with time throughout the duration of fatiguing isometric contractions at tensions between 25 and 100% MVC. During sustained maximal effort, the frequency initially decreased linearly with time. However, once the target tension could no longer be maintained, the centre frequency remained constant throughout the remainder of the contraction. The frequency was found to recover within 1 min following exercise at all tensions examined.

Adult

Inhibition in the release of immunoreactive beta-endorphin from the periaqueductal grey during isometric contractions of cat hind-limb muscles: the effects of clonidine.

Glass microelectrodes, coated with antibodies specific for beta-endorphin, were inserted into the right periaqueductal grey (PAG) (at PO.5-1.0 mm, LR 2.0 mm and 6.0 mm below the dorsal surface of the colliculi) of cats anesthetized with alpha-chloralose to determine whether immunoreactive beta-endorphins (ir-END) were released in response to fatiguing isometric contractions of the hind-limb muscles. Probes were inserted into the PAG prior to, during and following muscle contraction in the absence or presence of clonidine. ir-END was released from the PAG up to 3 h after surgery was completed while cats remained at rest. In the absence of clonidine, mean arterial pressure increased by 65 +/- 12 mmHg during contractions and the release of ir-END was inhibited during the contraction periods compared to resting periods. Clonidine abolished the pressor response to muscular contraction when injected into the PAG, but did not cause the release of ir-END during fatiguing isometric contractions. These data suggest that isometric contractions of skeletal muscle do not induce the release of ir-END-like substances from the PAG and clonidine does not attenuate the muscle pressor response by causing the release of ir-END from this level in the PAG.

Animals

Fatiguing isometric contraction of hind-limb muscles results in the release of immunoreactive neurokinins from sites in the rostral medulla in the anesthetized cat.

Antibody-coated microprobes were used to determine whether immunoreactive neurokinins (irNK) were released from sites in the brainstem during fatiguing isometric contractions of the triceps surae muscles in cats anesthetized with alpha-chloralose. Contractions were generated by stimulating the tibial nerve using a microprocessor-controlled stimulator. Microprobes were inserted into the periaqueductal grey (P 0.5-1.0 mm) or the medullary brainstem (either 3.0 or 3.5 mm rostral to the obex) prior to, during and following fatiguing contractions. No release of irNK was detected from the periaqueductal grey as a result of fatiguing isometric contractions. When probes were inserted 3.0 mm rostral to the obex, a basal release of irNK was detected from the medulla but this was inhibited during isometric contractions from sites corresponding to the lateral tegmental field. When probes were inserted into the more rostral site in the medulla (3.5 mm rostral to the obex), irNK were released in response to contractions from sites corresponding to lateral reticular nucleus, ventral regions of the nucleus tractus solitarius and the medial vestibular nucleus. No irNK were released from this site (3.5 mm rostral to obex) either during passive leg flexing, during nerve stimulation following gallamine injection and muscle paralysis or during stimulation of the central end of the cut tibial nerve. These results demonstrate that neurokinins are released from discrete sites in the medulla in response to fatiguing muscle contractions and suggest that tachykinin neurons may be a component of the pathways regulating blood pressure during ergoreceptor activation.

Anesthesia

Effects of carotid artery occlusion on the pressor response induced by sustained isometric contraction in the cat.

The effects of clonidine, a central alpha 2 agonist, on changes in blood pressure caused by muscle afferent nerve (ergoreceptor) activation and baroreceptor manipulation were studied in cats. Prolonged isometric contractions (ergoreceptor activation) of the gastrocnemius and plantaris muscles increased mean arterial pressure by 53 mmHg. This pressor response was not altered by naloxone (0.5 mumol.litre-1) but was eliminated by clonidine (0.5-2.0 micrograms) when injected into the cerebral aqueduct. Brief occlusion of the carotid artery (15-30 s) caused mean arterial pressure to increase by 32-42 mmHg at rest. Neither naloxone nor clonidine altered the magnitude of the reflex pressor response to carotid occlusion. Similar increases in pressure were measured when occlusion was applied during fatiguing isometric contractions; thus baroreceptor induced increases in pressure were superimposed on the ergoreceptor induced blood pressure changes. Naloxone did not affect the changes in pressure caused by either reflex response. Clonidine continued to eliminate the pressor response to muscular contraction but did not affect the pressure increase when the carotid occlusion was applied during contractions. Electrical stimulation of the carotid sinus nerve caused blood pressure to decrease by 36 mmHg during rest and by 41 mmHg during fatiguing isometric contractions. Clonidine did not alter the depressor response to carotid sinus nerve stimulation. These data may indicate that separate pathways centrally mediate the changes in blood pressure caused by ergoreceptor and baroreceptor afferent activation. The integration of the ergoreceptor pathway may involve a catecholaminergic-opioidergic system but the present results do not suggest a similar interaction for the baroreceptor integration.

Animals

The influence of location of electrode on muscle fiber conduction velocity and EMG power spectrum during voluntary isometric contraction measured with surface array electrodes.

A surface array electrode was used to investigate muscle fiber conduction velocity (MFCV) and EMG power spectrum during voluntary isometric contraction of m. biceps brachii. The mean power frequency (MPF) and the total power (TP) at various locations of the muscle were obtained from the power spectrum. MFCVs at various locations of the muscle were measured directly using the averaging method. The values of MPF, TP and MFCV were identified with respect to the electrode locations on the m. biceps brachii. MPF was shown as high near the end-plate and low near the tendon of the muscle during contraction of 40% of isometric maximum voluntary contraction (MVC). TP showed low value near the end-plate and the tendon of the muscle fiber and different values at different location setting electrode during voluntary isometric contraction. TP at each location on the muscle surface increased when the contraction levels increased in the contractions range of 20 to 60% MVC. MFCVs showed a high value near the end-plate and the tendon of the muscle during the contraction of 40% MVC. These results indicated that MPF, TP and MFCV at different locations on the muscle were different along the length of the muscle fiber.

Adult

Cardiovascular changes during an isometric contraction combined to a cold pressor test.

Fifteen normotensive male subjects (age, 27.9 +/- 1.3 years) performed isometric contractions (handgrip) at 15, 30 and 50% maximal voluntary contraction respectively for 7, 3 and 1 min. The contractions were sustained with or without a cold pressor test of the same duration (immersion of one foot in water of 5.2 +/- 0.1 degrees C). At rest, under the influence of a cold pressor test for 3 min, the heart rate increased from 74.3 +/- 2.2 to 83.3 +/- 2.1 beats min-1 (P < 0.001) in less than 10 s, and returned to the control value in 3 min; on the contrary, at the end of 3 min of cold pressor test, the systolic blood pressure was still above the control value (135.5 +/- 4.2 and 121.0 +/- 3.2 mmHg, P < 0.001). During the contractions, there was no significant difference between heart rate and systolic blood pressure values obtained with or without cold pressor test. Similarly, there was no significant change in the concentrations of plasma catecholamines. This absence of cardiovascular effect of cold pressor test associated with isometric contraction (for three relative force levels) is probably due to the increasing importance of sympathetic cardiac activation produced by the contraction with respect to that produced by the cold pressor test, the absence of supplementary changes in total peripheral resistance and a partial reciprocal inhibition of pain produced by the simultaneous performance of these two tests.

Adult

Effect of muscle mass on the pressor response in man during isometric contractions.

1. Changes in blood pressure and heart rate were measured in six healthy male subjects during voluntary isometric contractions of the forearm and quadriceps muscles. Arterial pressure was measured directly via a catheter inserted into the radial artery of the non-contracting arm. Each subject exerted two types of contractions: (a) sustained contractions at 70% of the maximum voluntary contraction (MVC) until fatigue occurred and (b) sustained contractions starting at maximum tension (100% MVC), held for a total duration of 1 min. 2. During fatiguing contractions at 70% MVC, there was a progressive increase in blood pressure, reaching a peak level at fatigue. The same level of mean arterial pressure was achieved during contractions of the same relative tension, regardless of the muscle mass. The same trend was observed for the changes in heart rate. 3. During contractions which started with the maximum tension, where tension fell continuously during the 60 s of maximal effort, mean arterial pressure rapidly increased to high levels within a few seconds, and then increased further by 20-30 mmHg during the sustained maximal effort. There was no difference in the initial rapid increases in mean arterial pressure, nor in the final mean arterial pressures reached between contractions of the forearm or quadriceps muscles. There were no differences in the heart rates achieved during these contractions either. 4. There was no significant difference between the mean arterial pressures observed at fatigue of a 70% MVC contraction or at the end of the 60 s maximum effort during handgrip contractions, or during contractions with the quadriceps muscles. 5. These results support the view that muscle mass is not a determinant of the magnitude of the cardiovascular reflexes during fatiguing isometric contractions in man.

Adult

Muscular pain caused by isometric contraction: evaluation of pain through visual analog scale.

Pain caused by muscular isometric contraction was investigated through use of the visual analog scale. The results demonstrate that the latency time and the time to reach maximum intensity are inversely proportional to the amount of the load; therefore increase of pain is linear. The high statistical significance of the results suggests that pain by isometric contraction could be a comparison test for somatic pain evaluation in spontaneous pathological conditions.

Adult

Rapid activation of glycogen synthase and protein phosphatase in human skeletal muscle after isometric contraction requires an intact circulation.

The effects of isometric contraction (66% of maximal force) and recovery on glycogen synthase fractional activity (GSF) in human skeletal muscle have been studied. Biopsies were taken from the quadriceps femoris muscle at rest, at fatigue and 5 min postexercise on two occasions: after one of the contractions, the circulation to the thigh was occluded during the 5 min recovery (OCC), and after the other contraction, the circulation was intact (control, CON). During CON, GSF decreased from (mean +/- SE) 0.34 +/- 0.05 at rest to 0.24 +/- 0.02 at fatigue and then increased to 0.74 +/- 0.04 at 5 min postexercise; corresponding values for OCC were 0.37 +/- 0.04, 0.25 +/- 0.04 and 0.48 +/- 0.05 (P < 0.001 vs. CON for 5 min postexercise only). Compared with the value at fatigue, protein phosphatase activity (PP) increased by 79 +/- 16% during CON recovery (P < 0.01), whereas no change was observed during OCC recovery. Uridine diphosphate glucose increased by approximately 2.5-fold at fatigue, remained elevated during OCC recovery, but reverted to the preexercise level during CON recovery (P < 0.001 vs. OCC recovery). Glucose 6-P increased approximately 5-fold at fatigue and was higher at 5 min postexercise in OCC vs. CON recovery (8.6 +/- 1.5 vs. 4.1 +/- 0.9 mmol/kg dry wt; P < 0.01). It is concluded that the rapid increase in GSF after intense exercise with an intact circulation may be at least partly attributed to an increase in the specific activity of PP. The increase in GSF during recovery in OCC may be at least partly attributed to the high glucose 6-P content in vivo, which enhances the substrate suitability of GS for PP. Thus, separate mechanisms exist for the activation of PP and GS during recovery from intense short term exercise.

Adult

Identification of time-varying dynamics of the human triceps surae stretch reflex. I. Rapid isometric contraction.

We have examined the time variations of stretch reflex dynamics throughout rapid voluntary changes in the isometric contraction level of the human triceps surae muscles. This was achieved by superimposing a small stochastic displacement upon many such changing contractions and then identifying the time-varying relationship between the perturbation and the evoked electromyograms (EMGs). An "ensemble" time-varying system identification technique was used to estimate these input-output dynamics as a set of impulse response functions, one for each time before, during, and after the change in contraction level, with a temporal resolution equal to the data acquisition rate. Three main findings resulted. First, stretch reflex gain (relating joint velocity to EMG) was significantly modulated during changes in voluntary contraction level, increasing as the subject contracted the muscles and decreasing as the subject relaxed. Second, stretch reflex dynamics did not change with contraction level, even when its gain varied substantially. Third, the time course of the gain changes closely followed the level of the EMG, even though the subjects used rather different activation and deactivation patterns. These results suggest that, for the behavior studied (i.e., rapid changes in isometric contraction level), stretch reflex gain and motoneuron pool activation level were controlled by a common descending command rather than being independently specified.

Adult

Effects of hypoxemia with and without acidemia on the isometric contraction time and the electromechanical delay of the fetal myocardium: an experimental study on the ovine fetus.

This study assessed the response of the preejection period during hypoxemia with and without acidemia. In five pregnant ewes, hypoxemia was created during 1 hour followed by fetal infusion of lactic acid during 2 hours. A micromanometer catheter positioned above the fetal aortic valve, an endocavitary phonocardiogram, and a fetal electrocardiogram allowed measurements of the two components of the preejection period--the isometric contraction time and the electromechanical delay. At the onset of hypoxemia, because of changes in isometric contraction time, the preejection period began to shorten. When acidemia was induced, the preejection period modified slowly in the opposite direction and lengthened, initially because of a prolongation of electromechanical delay and later because of an increase in the already shortened isometric contraction time. This process developed slowly and at the end of 2 hours of acidemia, preejection period were back to preexperimental values. It can be concluded that systolic time intervals can be normal and misleading when acidosis complicates hypoxemia.

Acids

Motor unit recruitment in human medial gastrocnemius muscle during combined knee flexion and plantarflexion isometric contractions.

Previous work on multifunctional muscle has suggested that motor unit recruitment during a combined force task is the result of an interactive effect of weighted inputs acting simultaneously on the motoneuron pool. The present study shows that a similar effect describes motor unit activation in a two-joint muscle as forces are combined at both proximal and distal attachments. The recruitment thresholds of single motor units in medial gastrocnemius muscle were determined during combined knee flexion and plantarflexion isometric contractions. Slow isometric ramp contractions in knee flexion were produced while maintaining various background levels of plantarflexion force. The combination of knee flexion and plantarflexion forces at which a motor unit initially discharged was used to characterize recruitment as represented by the slope of the regression line fit to the individual data points. Each subject completed two experiments; one at each of two knee joint angles, with the ankle joint fixed at 90 degrees. The effect of knee angle was assessed by comparing the slopes of the regression lines that characterized motor unit recruitment at each knee angle. Motor units in medial gastrocnemius were recruited when the linear sum of the forces exerted in plantarflexion and knee flexion exceeded a certain threshold of combined force. Specifically, the apparent force threshold of recruitment in knee flexion decreased as the level of force maintained in plantarflexion increased. Further, evidence is provided indicating that the linear relationship describing recruitment in two-joint muscle is dependent upon joint angle.(ABSTRACT TRUNCATED AT 250 WORDS)

Adult

The effect of forskolin on the isometric contraction of the isolated hemidiaphragm of the rat.

The effects of forskolin on the parameters of the isometric contraction alone and in combination with aminophylline and isoprenaline were studied on the isolated hemidiaphragm of the rat during direct electrical stimulation. Forskolin (2.6 - 18.2 mumol l-1) produced a concentration-dependent increase in tension developed (Td) and, to a lesser extent, in the maximum rate of rise in tension (dT/dt max). The dose-response curve for the action of forskolin (2.6 - 18.2 mumol l-1) on Td was shifted to the left in the presence of a standard concentration of aminophylline (0.32 mmol l-1). Forskolin (5.2 mumol l-1) produced a further and significant increase in both Td and dT/dt max in the presence of isoprenaline (0.24 mumol l-1) in the bath. In a calcium-free medium, the effects of forskolin (7.80 and 18.2 mumol l-1) on Td and dT/dt max were significantly weaker than in a medium containing calcium. These data indicate that forskolin increases the isometric contraction of the isolated hemidiaphragm probably by activating the adenosine 3':5'-cyclic monophosphate (cyclic AMP) generating system. These effects are possible only in the presence of calcium.

Aminophylline

Energy metabolism in different human skeletal muscles during voluntary isometric contractions.

The energy turnover in contracting skeletal muscle was studied by measuring the rate of temperature rise during voluntary, isometric contractions and circulatory arrest in M. soleus, M. sacrospinalis and M. biceps brachii in 14 males, by thermoelements inserted in the muscles. A linear relationship between rate of temperature rise and force intensity given as per cent of maximal voluntary contraction (MVC) was demonstrated in biceps (r = 0.95), but not so clearly confirmed in soleus (r = 0.73). Muscle biopsies were taken from the same muscles and fibre type distribution was determined histochemically by staining for ATPase. The rate of heat production at MVC showed positive correlation to the percentage of fast twitch (FT) fibres in the muscles (r = 0.90). Linear extrapolation indicates that the maximal energy turnover in human FT fibres is approximately six times that of slow twitch (ST) fibres during voluntary isometric contractions.

Adult

Muscle ATP turnover rate during isometric contraction in humans.

ATP turnover and glycolytic rates during isometric contraction in humans have been investigated. Subjects contracted the knee extensor muscles at two-thirds maximal voluntary force to fatigue (mean +/- SE, 53 +/- 4 s). Biopsies were obtained before and after exercise and analyzed for high-energy phosphates and glycogenolytic-glycolytic intermediates. Total ATP turnover was 190 +/- 7 mmol/kg dry muscle, whereas the average turnover rate was 3.7 +/- 0.2 mmol . kg dry muscle-1 . S-1. The average ATP turnover rate was positively correlated with the percentage of fast-twitch fibers in the postexercise biopsy (r = 0.71; P less than 0.05) and negatively correlated with contraction duration to fatigue (r = -0.88; P less than 0.05). At fatigue, phosphocreatine ranged from 1 to 11 mmol/kg dry muscle (86-99% depletion of value at rest), whereas lactate ranged from 59 to 101. The mean glycolytic rate was 0.83 +/- 0.05 mmol . kg dry muscle-1 . S-1 and was positively correlated with the rate of glucose 6-phosphate accumulation (r = 0.83; P less than 0.05). It is concluded that a major determinant of the ATP turnover rate is the muscle fiber composition, which is probably explained by a higher turnover rate in fast-twitch fibers; fatigue is more closely related to a low phosphocreatine content than to a high lactate content; and the increase in prephosphofructokinase intermediates is important for stimulating glycolysis during contraction.

Adenosine Triphosphate