Central mechanisms of fatigue during local static muscular activity

Changes in blood circulation, heart electrical activity, and the function of the brain cortex have been studied in healthy subjects of six age groups (from 7 to 65 years of age) during local static exercise. It has been shown that systemic responses of the body, such as a decrease in the contractility of muscles, changes in the heart function due to enhancement of central sympathetic regulation, and changes in the type of brain bioelectrical activity recorded with the use of encephalography, develop during fatigue.     

Thin-fiber mechanoreceptors reflexly increase renal sympathetic nerve activity during static contraction

The renal vasoconstriction induced by the sympathetic outflow during exercise serves to direct blood flow from the kidney toward the exercising muscles. The renal circulation seems to be particularly important in this regard, because it receives a substantial part of the cardiac output, which in resting humans has been estimated to be 20%. The role of group III mechanoreceptors in causing the reflex renal sympathetic response to static contraction remains an open question. To shed some light on this question, we recorded the renal sympathetic nerve responses to static contraction before and after injection of gadolinium into the arterial supply of the statically contracting triceps surae muscles of decerebrate unanesthetized and chloralose-anesthetized cats. Gadolinium has been shown to be a selective blocker of mechanogated channels in thin-fiber muscle afferents, which comprise the afferent arm of the exercise pressor reflex arc. In decerebrate (n = 15) and chloralose-anesthetized (n = 12) cats, we found that gadolinium (10 mM; 1 ml) significantly attenuated the renal sympathetic nerve and pressor responses to static contraction (60 s) after a latent period of 60 min; both responses recovered after a latent period of 120 min. We conclude that thin-fiber mechanoreceptors supplying contracting muscle are involved in some of the renal vasoconstriction evoked by the exercise pressor reflex.     

Different responses in skin and muscle sympathetic nerve activity to static muscle contraction

We microneurographically recorded the traffic of sympathetic nerves leading to foot volar skin activity (SSA) and leg skeletal muscle activity (MSA) during isometric handgrip and simultaneously determined sweat rate by the ventilated capsule method and skin blood flow by laser-Doppler flowmetry in the innervating area of SSA. SSA increased abruptly and was almost constant during handgrip, accompanied by an increase in sweat rate, whereas skin blood flow showed no significant change during the handgrip. MSA showed a time-dependent increase during the course of handgrip. During arterial occlusion of the working forearm after handgrip, SSA decayed to the precontraction control level, whereas MSA remained at a higher level than during control. During involuntary biceps muscle contraction induced by electrical stimulation, both SSA and MSA increased. The results suggest that the SSA response during voluntary handgrip, which was demonstrated to contain mainly sudomotor activity, might be influenced by central command and input from peripheral mechanoreceptors but be influenced little by input from muscle chemoreceptors.     

Sympathetic nerve responses to muscle contraction and stretch in ischemic heart failure

Congestive heart failure (CHF) induces abnormal regulation of peripheral blood flow during exercise. Previous studies have suggested that a reflex from contracting muscle is disordered in this disease. However, there has been very little investigation of the muscle reflex regulating sympathetic outflows in CHF. Myocardial infarction (MI) was induced by the coronary artery ligation in rats. Echocardiography was performed to determine fractional shortening (FS), an index of the left ventricular function. We examined renal and lumbar sympathetic nerve activities (RSNA and LSNA, respectively) during 1-min repetitive (1- to 4-s stimulation to relaxation) contraction or stretch of the triceps surae muscles. During these interventions, the RSNA and LSNA responded synchronously as tension was developed. The RSNA and LSNA responses to contraction were significantly greater in MI rats (n = 13) with FS <30% than in control animals (n = 13) with FS >40% (RSNA: +49 +/- 7 vs. +19 +/- 4 a.u., P < 0.01; LSNA: +28 +/- 7 vs. +8 +/- 2 a.u., P < 0.01) at the same tension development. Stretch also increased the RSNA and LSNA to a larger degree in MI (n = 13) than in control animals (n = 13) (RSNA: +36 +/- 6 vs. +19 +/- 3 a.u., P < 0.05; LSNA: +24 +/- 3 vs. +9 +/- 2 a.u., P < 0.01). The data demonstrate that CHF exaggerates sympathetic nerve responses to muscle contraction as well as stretch. We suggest that muscle afferent-mediated sympathetic outflows contribute to the abnormal regulation of peripheral blood flow seen during exercise in CHF.     

Development of muscle fatigue during intermittent submaximal static contraction in an agonist heterogeneous muscle group.

A comparison of the mean power frequency (MPF) and the root mean square amplitude (rms) of the myo-electric signal of two agonist muscles [triceps brachii (fast; TB) and anconeus (slow; ANC)] has been made during repeated intermittent static contractions. Subjects were asked to maintain different extension torques at 50% of maximal voluntary contraction until this could no longer be maintained (endurance time). The interval between successive contractions was kept constant at 3 min. During the first six successive contractions, a decrease in MPF and an increase in rms were most pronounced, ANC and TB MPF recovered with subsequent overshoot. A marked decline in endurance time was also seen. The increase in rms was greater for TB than for ANC when the decrease in MPF was greater for ANC than for TB. The differences in power spectrum density function upper frequencies of the two muscles could explain the greater decrease of MPF in ANC. Our data would suggest a greater fatigability in TB relative to ANC. On and after the seventh contraction, a steady-state in duration, muscle temperature, MPF and rms was reached. These results suggested that a slow (ANC) and a fast (TB) muscle acted in a similar way during intermittent static contractions, when the intervening rest was not long enough to allow full recovery of the muscles.     

Thermal sensation and comfort during exposure to local airflow to face or legs

The present study examined the contribution of local airflow temperature to thermal sensation and comfort in humans. Eight healthy male students were exposed to local airflow to their faces (summer condition) or legs (winter condition) for 30 minutes. Local airflow temperature (Tf) was maintained at 18 degrees C to 36 degrees C, and ambient temperature (Ta) was maintained at 17.4 degrees C to 31.4 degrees C. Each subject was exposed to 16 conditions chosen from the combination of Tf and Ta. Based on the results of multiple regression analysis, the standardized partial regression coefficient of Tf and Ta were determined to be 0.93 and 0.13 in the summer condition, and 0.71 and 0.36 in the winter condition at the end of the exposure. Also, thermal comfort was observed to depend closely on the interrelation between Tf and Ta. The present data suggested that local airflow temperature is an important thermal factor regarding thermal sensation and comfort.     

Cardiovascular responses to static and dynamic contraction during comparable workloads in humans

Previous studies suggest that the blood pressure response to static contraction is greater than that caused by dynamic exercise. In anesthetized cats, however, pressor responses to electrically induced static and dynamic contraction of the same muscle group are similar during equivalent workloads and peak tension development [i.e., similar tension-time index (TTI)]. To determine if the same relationship exists in humans, where contraction is voluntary and central command is present, dynamic (180 s; 1/s) and static (90 s) contractions at 30% of maximal voluntary contraction (MVC) were performed. Dynamic contraction also was repeated at the same TTI for 90 s at 60% MVC. Mean arterial pressure (MAP), heart rate (HR), cardiac output (CO), MAP during postexercise arterial occlusion (an index of the metaboreceptor-induced activation of the exercise pressor reflex), and relative perceived exertion (RPE) (an index of central command) were assessed. No differences in these variables were found between static and dynamic contraction at a tension of 30% MVC. During dynamic contraction at 60% MVC, changes in MAP (16 +/- 3 vs. 19 +/- 4 mmHg) and absolute HR (92 +/- 6 vs. 69 +/- 5 beats/min), CO (7.9 +/- 0.4 vs. 6.3 +/- 0.3 l/min), RPE (16 +/- 1 vs. 13 +/- 1), and MAP during postexercise arterial occlusion (115 +/- 3 vs. 100 +/- 4 mmHg) were greater than during static contraction (P < 0.05). Thus increases in MAP and HR, activation of central command, and muscle metabolite-induced stimulation of the exercise pressor reflex during static and dynamic contraction in humans seem to be similar when peak tension and TTI are equal. Augmented responses to dynamic contraction at 60% MVC are likely related to greater activation of these two mechanisms.     

Sympathetic nerve activity in arm and leg muscles during lower body negative pressure in humans

Nonhypotensive lower body negative pressure (LBNP) is reported to decrease forearm but not calf blood flow as measured by strain-gauge plethysmography. This suggests that unloading of cardiopulmonary receptors increases sympathetic outflow to arm but not to leg. To test this hypothesis we measured muscle sympathetic nerve activity (MSA) in the arm (radial nerve) and leg (peroneal nerve) simultaneously during LBNP. In eight healthy subjects, we measured heart rate, blood pressure, and radial and peroneal MSA during LBNP at 10 and 20 mmHg. There was no difference between radial and peroneal MSA at rest, and there were successive parallel increases of MSA in both nerves during LBNP at 10 and 20 mmHg. These data indicate that there are nearly identical increases of sympathetic outflow to the arm and leg during mild to moderate degrees of orthostatic stress.     

Non-uniform mechanical activity of quadriceps muscle during fatigue by repeated maximal voluntary contraction in humans

To determine the non-uniform surface mechanical activity of human quadriceps muscle during fatiguing activity, surface mechanomyogram (MMG), or muscle sound, and surface electromyogram (EMG) were recorded from the rectus femoris (RF), vastus lateralis (VL), and vastus medialis (VM) muscles of seven subjects during unilateral isometric knee extension exercise. Time- and frequency-domain analyses of MMG and of EMG fatigued by 50 repeated maximal voluntary contractions (MVC) for 3 s, with 3-s relaxation in between, were compared among the muscles. The mean MVC force fell to 49.5 (SEM 2.0)% at the end of the repeated MVC. Integrated EMG decreased in a similar manner in each muscle head, but a marked non-uniformity was found for the decline in integrated MMG (iMMG). The fall in iMMG was most prominent for RF, followed by VM and VL. Moreover, the median frequency of MMG and the relative decrease in that of EMG in RF were significantly greater (P < 0.05) than those recorded for VL and VM. These results would suggest a divergence of mechanical activity within the quadriceps muscle during fatiguing activity by repeated MVC. Accepted: 19 January 1999     

Respiratory sensation and pattern of respiratory muscle activation during diaphragm fatigue

We have examined the relationship between respiratory effort sensation (modified Borg scale) and amplitude of the integrated surface electromyogram of the diaphragm (Edi, esophageal electrode), rib cage muscles (Erc), and sternomastoid muscle (Esm) during the development of diaphragm fatigue in five normal subjects. Three conditions were studied: run A: transdiaphragmatic pressure (Pdi), 65% Pdimax; esophageal pressure (Pes), 60% Pesmax; run B: Pdi, 50% Pdimax; Pes, 60% Pesmax; and run C: Pdi, 50% Pdimax; Pes, 20% Pesmax. During all runs there was a progressive rise in sensation, which was greater in runs A and B than in run C (P less than 0.05, analysis of variance). There was no difference between runs A and B. At the end of run C subjects did not report a maximal Borg score despite their inability to generate the target Pdi. The increase in sensory score with fatigue correlated highly with Esm/Esmmax and with Erc/Ercmax. There was no correlation between sensory score and Edi/Edimax. We conclude that the increase in respiratory effort sensation that accompanies diaphragm fatigue is not due to perception of increased diaphragmatic activation. It may reflect increased overall respiratory motor output not directed to the diaphragm.     

Sympathetic nerve activity and whole body heat stress in humans

We and others have shown that moderate passive whole body heating (i.e., increased internal temperature ～0.7°C) increases muscle (MSNA) and skin sympathetic nerve activity (SSNA). It is unknown, however, if MSNA and/or SSNA continue to increase with more severe passive whole body heating or whether these responses plateau following moderate heating. The aim of this investigation was to test the hypothesis that MSNA and SSNA continue to increase from a moderate to a more severe heat stress. Thirteen subjects, dressed in a water-perfused suit, underwent at least one passive heat stress that increased internal temperature ～1.3°C, while either MSNA (n = 8) or SSNA (n = 8) was continuously recorded. Heat stress significantly increased mean skin temperature (Δ～5°C, P < 0.001), internal temperature (Δ～1.3°C, P < 0.001), mean body temperature (Δ～2.0°C, P < 0.001), heart rate (Δ～40 beats/min, P < 0.001), and cutaneous vascular conductance [Δ～1.1 arbitrary units (AU)/mmHg, P < 0.001]. Mean arterial blood pressure was well maintained (P = 0.52). Relative to baseline, MSNA increased midway through heat stress (Δ core temperature 0.63 ± 0.01°C) when expressed as burst frequency (26 ± 14 to 45 ± 16 bursts/min, P = 0.001), burst incidence (39 ± 13 to 48 ± 14 bursts/100 cardiac cyles, P = 0.03), or total activity (317 ± 170 to 489 ± 150 units/min, P = 0.02) and continued to increase until the end of heat stress (burst frequency: 61 ± 15 bursts/min, P = 0.01; burst incidence: 56 ± 11 bursts/100 cardiac cyles, P = 0.04; total activity: 648 ± 158 units/min, P = 0.01) relative to the mid-heating stage. Similarly, SSNA (total activity) increased midway through the heat stress (normothermia; 1,486 ± 472 to mid heat stress 6,467 ± 5,256 units/min, P = 0.03) and continued to increase until the end of heat stress (11,217 ± 6,684 units/min, P = 0.002 vs. mid-heat stress). These results indicate that both MSNA and SSNA continue to increase as internal temperature is elevated above previously reported values.     

Changes in muscle sympathetic nerve activity and calf blood flow during static handgrip exercise

To test the function of sympathetic vasco-constrictor nerves on blood flow in resting limbs during static muscle contraction, muscle sympathetic nerve activity (MSNA) to the leg muscle was recorded from the tibial nerve microneurographically before, during and after 2 min of static handgrip (SHG). Simultaneously, calf blood flow (CBF) was measured by strain gauge plethysmography. An increase in MSNA, a decrease in CBF and an increase in calf vascular resistance (CVR) in the same resting limb occurred concomitantly during SHG. However, the increase in CVR was blunted in the second minute of handgrip when MSNA was still increasing. The results indicated that the decrease of CBF during SHG reflects the increase in MSNA, while the dissociation between MSNA and CVR at the later period of SHG may be related to metabolic change produced by the vasoconstriction.     

Electromyography and fatigue during prolonged, low-level static contractions

Findings from five separate studies of EMG changes and muscle fatigue during prolonged low-level static contractions are summarized, and the possible mechanisms behind the changes are briefly discussed. Sustained static contractions (10%, 7% and 5% MVC) of up to 1 h duration were performed by finger flexors, elbow flexors and extensors, and knee extensors. In one experiment, intermittent static arm pulling (triceps) (10 s contraction and 5 s rest, average work load 14% and 10% MVC) was performed for 7 h. The endurance time for the sustained contractions was around one hour for 10% MVC, and it was shown--all in all--that the concept of "indefinite" endurance times at contractions below 15-20% MVC cannot be maintained. After 5% MVC sustained contractions for one hour a 12% reduction in MVC was seen, and significant increases in EMG amplitude and decreases in the mean spectral frequency of the EMG-power spectrum were found. Marked differences were also seen in the EMG changes in the elbow flexors and extensors, and transcutaneous electrical stimulation of the knee extensors showed that low frequency fatigue was present after the contraction. With intermittent contractions similar changes in the EMG parameters were seen after 2-3 h of contractions at 14% MVC. On average, during contractions of 10% MVC no EMG changes were detected. Increased extracellular potassium concentration in the contracting muscles is suggested as a possible explanation of these findings.     

Physiological and psychological indices of fatigue during static contractions

The development of fatigue and the relationship between psychological and physiological indices of fatigue were studied in a group of 18 male subjects during static contractions. Exercise was performed as a static elbow flexion at 25% MVC. Heart rate (HR), intraarterial blood pressure (BP) and surface EMG [mean amplitude (MA) and central frequency (CF)] were studied during contractions sustained until exhaustion. The amount of effort expended (relative to total exhaustion) and the rating of perceived pain were recorded following contractions interrupted after 20, 30 . . . 80, and 100% of endurance time. HR, BP and EMG amplitude responses were similar to those previously recorded. The decline in CF occurred in two phases, possibly related to a change in motor unit recruitment after the initial 70% of endurance time. The subjects overestimated the amount of effort expended and thus underestimated their endurance capacity. The best correlation between perceived effort and physiological responses was obtained using blood pressure data, whereas changes in EMG data did not parallel the psychological responses. It is concluded that the perception of effort during a static contraction is produced through a complex process, in which several influences of peripheral and central origin are integrated.