Exhausting stretch-shortening cycle (SSC) exercise causes greater impairment in SSC performance than in pure concentric performance

The purpose of the present study was to investigate the fatigue effect of repeated exhaustive stretch-shortening cycle (SSC) exercise on concentric muscle function. Ten healthy male subjects performed SSC exercise [92 (30) jumps] on a special sledge apparatus. Exhaustion occurred on average within 3 min. A squat jump (SJ) test utilizing a concentric-only action was performed immediately before and after the SSC exercise, and then 10 min, 20 min, 2 days and 4 days later. In addition, a drop jump (DJ) test using an SSC was also performed immediately before and 20 min after the SSC exercise, and 2 days and 4 days later. During jump tests, lower limb joint moment, power, and work contributions were analyzed by using the kinetic and kinematic data. The fatigue exercise was characterized by a relatively high blood lactate concentration [7.2 (0.8) mmol·l^–1] and a 2-day delayed increase in serum creatine kinase activity [486 (300) U·l^–1]. SJ performance decreased markedly immediately after the SSC exercise (P<0.05) and then recovered within 10 min. In contrast, DJ performance and knee joint contribution showed a delayed decrease 2 days after the SSC exercise bout. The surface electromyographic (EMG) activity of the lower limb muscles showed no obvious change in the SJ in comparison to the DJ, although in the latter there was a delayed decrease of knee extensor EMG during the pre-activation and braking phases. The results suggest that isolated concentric muscle function is affected mainly by acute metabolic fatigue after SSC exercise. During a follow-up period after the exercise, changes in hip and knee joint contribution in SJ showed a different recovery pattern compared to those in eccentric DJ. It could be suggested that exhaustive SSC exercise would mainly influence the relative power–work balance between the hip and knee joints during the eccentric phase of SSC. Thus different motor control strategies may account for the distinctive fatigue responses observed in SJ and DJ. Electronic Publication     

Reduced stretch-reflex sensitivity after exhausting stretch-shortening cycle exercise

The stretch-shortening cycle (SSC) is an effective and natural form of muscle function but, when repeated with sufficient intensity or duration, it may lead to muscle damage and functional defects. A reduced tolerance to impact has been reported, which may be partly attributed to a reduced stretch-reflex potentiation. The aim of the present study was to examine the influence of SSC-induced metabolic fatigue and muscle damage on the efficacy of stretch reflexes, as judged by the electromyograph (EMG) response of two shank muscles (lateral gastrocnemius LG, soleus SOL) to controlled ramp stretches. These EMG responses were recorded before and immediately after exhausting SSC-type leg exercise and 2 h, 2 days and 4 days later. Serum concentrations of creatine kinase ([CK]), myoglobin and lactate were measured repetitively along the protocol. Two maximal vertical drop jumps and counter-movement jumps were performed after each reflex test. The exhausting SSC-type exercise induced an immediate reduction (P < 0.05) with a delayed short-term recovery of the LG peak-to-peak reflex amplitude. This was not accompanied by significant changes in the reflex latency. The drop jump performance remained slightly but significantly reduced (P < 0.05) until the 2nd day postexercise. Peak [CK] appeared for all the subjects on the 2nd day, suggesting the presence of muscle damage. The increase in [CK] between the 2nd h and the 2nd day postexercise was found to be negatively related (P < 0.001) to the relative changes in the drop jump height. Furthermore, a significant relationship (P < 0.05) was found between recovery of the stretch reflex in LG and the decrease of [CK] between the 2nd and the 4th day. hese findings support the hypothesis of a reduced stretch-reflex sensitivity. While the exact mechanisms of the reflex inhibition remain unclear, it is emphasized that the delayed recovery of the reflex sensitivity could have resulted from the progressive inflammation that develops in cases of muscle damage.     

Effects of long- and short-term fatiguing stretch-shortening cycle exercises on reflex EMG and force of the tendon-muscle complex

This study examined the fatigue effects of stretch-shortening cycle exercises of different intensity and duration on stretch reflex EMG and mechanical responses of the triceps surae muscle. Twelve subjects performed either a 10-km run (n=6) or short but exhaustive rebound exercise on a sledge apparatus (n=6). Passive reflex tests (mechanically induced ankle dorsiflexions) were examined before, after as well as 2 h, 2 and 7 days after exercise. Mechanical reflex responses were recorded from the ergometer torque signal. An acute contractile failure was observed as large reductions in twitch responses, especially in the sledge subgroup who showed high post-exercise peak blood lactate and an increased EMG/torque ratio. Independently of the exercise, the delayed fatigue analysis revealed strong relationships between the reflex-induced EMG and mechanical changes. In addition to muscle damage, these results may be explained by inhibitory effects via the sensitisation of small muscle afferents particularly during the exercise-induced delayed recovery process.     

Changes in the soleus muscle architecture after exhausting stretch-shortening cycle exercise in humans

This study focused on the architectural changes in the muscle–tendon complex during the immediate and secondary (delayed) reductions of performance (bimodal recovery) caused by an exhaustive rebound type stretch-shortening cycle (SSC) exercise. The isometric plantar flexor torque during maximum voluntary contraction (MVC) was measured together with recording of electromyography (EMG) and ultrasonography from the soleus muscle before (BEF), after (AFT), 2 h (2H), 2 and 8 days (2D, 8D) after the SSC exercise (n = 8). The performance variables (MVC torque and EMG activation) followed the bimodal recovery patterns. This was not the case in the changes of the fascicle length and muscle thickness. The relative torque changes in MVC correlated positively (R = 0.78, P = 0.02) to the corresponding averaged EMG changes between BEF and 2H (BEF → 2H); the significance disappeared in the comparison between 2H and 2D (2H → 2D), during which period MVC showed a secondary reduction. The relative torque changes in MVC showed no correlation with the changes in muscle thickness between BEF–2H. However, this correlation between 2H–2D was negative (R = –0.85, P < 0.01). The fascicle shortening/average EMG ratio in MVC increased at 2H, and then decreased more at 2D than 2H (P < 0.05). Thus, the secondary performance decline was not related to the corresponding EMG reduction but to the increased muscle thickness, which peaked at 2D. The results suggest clearly that the secondary decline in MVC could be related to the increase in muscle volume.     

Voluntary activation and mechanical performance of human triceps surae muscle after exhaustive stretch-shortening cycle jumping exercise

The purpose of this study was to examine neuromuscular factors that may contribute to post exercise force loss and subsequent recovery after exhaustive stretch-shortening cycle (SSC) exercise. Six subjects were fatigued on a sledge apparatus by 100 maximal rebound jumps followed by continuous submaximal jumping until complete exhaustion. Exercise-induced changes in neuromuscular performance were followed up to 7 days post exercise. The total number of jumps in the SSC exercise ranged from 336 to 1392. The SSC exercise induced a significant immediate plantarflexion torque decline of 29, 38 and 44% (P<0.05) in maximal voluntary contraction and evoked maximal twitch and low-frequency (LF) stimulation, respectively. The higher the number of jumps in the SSC exercise the larger was the post exercise reduction in voluntary activation as well as in contractile force (r=–0.94, P<0.01, in both). Furthermore, a higher number of jumps augmented a delayed force recovery and late decline in stretch reflex EMG response (r=–0.94, P<0.01). Clear differences were found in central and peripheral adaptation to the exhaustive SSC exercise between the subjects. The magnitude of post exercise contractile and activation failure as well as the delayed recovery of neuromuscular performance may have been augmented in some subjects due to their high number of jumps in the exercise.     

The influence of agonist premotor silence and the stretch-shortening cycle on contractile rate in active skeletal muscle

Summary Agonist premotor silence (PMS), a brief period of relative quiescence in active skeletal muscle prior to phasic activation, was investigated in subjects performing maximal contractions. The frequency of occurrence and potential function of the silent period were examined for elbow flexions and extensions. PMS was evident for movements in both directions, indicating that the mechanism is not primarily limited to extensors as previously hypothesized. Flexions demonstrating PMS exhibited increased velocity and acceleration; however, kinematic facilitation was only evident on trials exhibiting the muscular stretch-shortening cycle (SSC). The SSC was present on trials lacking PMS, demonstrating that biceps and triceps silence are not the sole determinants of preparatory agonist lengthening for elbow flexions and extensions, respectively. Taken together, the data indicate that agonist PMS is a mechanism under apparent central control that acts concomitantly with mechanical factors to potentiate elbow flexor contractions.     

Contraction-specific differences in maximal muscle power during stretch-shortening cycle movements in elderly males and females

Elderly people (age 75 years; n=48 males and 34 females) were studied in order to elucidate gender differences in elderly subjects on the determinants of muscle power (force and velocity) during a stretch-shortening cycle. All subjects performed three maximal counter-movement vertical jumps using both legs, on a force platform (Kistler 9281 B). The eccentric (Ep) and concentric (Cp) phases of the jumps were analyzed. The Ep was further divided into an acceleration phase (Ep_acc: from the start of the downward movement to the maximal negative velocity) and deceleration phase (Ep_dec: from the maximal negative velocity to the end of the downward movement). Jump height for the men was higher than for the women (P < 0.001). During both Ep_acc and Ep_dec no significant differences were observed between males and females in force and power generation. However, the men had a higher peak muscle power during the Cp, which may be explained exclusively by the velocity determinant (P < 0.001). No specific gender-related strategy appeared to influence the motor pattern of the movement. The comparable eccentric force generation of the leg extensors in both genders suggests a similar ability to cope with eccentric muscle actions during everyday activities. In contrast, the marked lower capacity for concentric contractions in women may result in an impaired performance, especially in activities where intense and rapid movements are essential, for example when reversing a forward fall. This may be one reason why elderly women are more prone to falls than are elderly men. Accepted: 19 September 2000     

Reduced stretch reflex sensitivity and muscle stiffness after long-lasting stretch-shortening cycle exercise in humans

It has been suggested that during repeated long-term stretch-shortening cycle (SSC) exercise the decreased neuromuscular function may result partly from alterations in stiffness regulation. Therefore, interaction between the short latency stretch-reflex component (M₁) and muscle stiffness and their influences on muscle performance were investigated before and after long lasting SSC exercise. The test protocol included various jumps on a sledge ergometer. The interpretation of the sensitivity of the reflex was based on the measurements of the patellar reflexes and the M₁ reflex components. The peak muscle stiffness was measured indirectly and calculated as a coefficient of the changes in the Achilles tendon force and the muscle length. The fatigue protocol induced a marked impairment of the neuromuscular function in maximal SSC jumps. This was demonstrated by a 14.1%–17.7% (n.s. –P?P?1 area under the electromyograms. Both of these methods of assessing the short latency reflex response showed a clear deterioration in the sensitivity of the reflex after fatigue (P?P?相似文献   

Stretch shortening cycle fatigue: interactions among joint stiness,reflex, and muscle mechanical performance in the drop jump

The purpose of the present study was to investigate the effect of strenuous stretch-shortening cycle exercise on the relationship between reflex and stiffness regulation during the drop jump. Ten healthy male subjects performed submaximal stretch-shortening cycle exercise on a special sledge apparatus. Exhaustion occurred on average within 3 min. A drop jump test from a 50-cm height was performed immediately before and after the sledge exercise, as well as 2 h, 2 days and 4 days later. The fatigue exercise showed relatively high blood lactate concentrations 12.5 (SD 2.6) mmol·l^–1 and a 2-day delayed increase of serum cretaine kinase concentration. In drop jumps, the short latency M1 component of the vastus lateralis muscle electromyogram (EMG) response showed a continuous decline throughout the entire follow-up period after fatigue (NS), whereas the medium latency EMG component increased 2 days after the postfatigue sessions (P < 0.05). Immediately after the fatigue exercise a positive correlation (P < 0.05) was found between the changes in the short latency EMG response and in the amount of knee joint stiffness during the early post-landing phase of the drop jump. This suggests that the M1 response was closely related to the stiffness changes during the initial braking phase of the drop jump. Increase of creatine kinase concentration on the 2nd day correlated negatively with the changes in the drop jump performance (P < 0.05). Since the short latency EMG component has almost recovered on the 2nd day, impairment of the mechanical function of the muscle might have taken place. In conclusion, exhausting stretch-shortening cycle exercise induced local muscle impairment, which resulted in modulation of the reflex and stiffness interaction in the drop jump as well as compensation by central motor command.     

Interaction between pre-landing activities and stiffness regulation of the knee joint musculoskeletal system in the drop jump: implications to performance

The purpose of the present study was to investigate the interaction between the pre-landing activities and the stiffness regulation of the knee joint musculoskeletal system and the takeoff speed during a drop jump (DJ). Nine healthy male subjects performed a DJ test from the height of 50 cm. The surface electromyographic (EMG) activity of the vastus lateralis (VL) muscle was recorded to evaluate both the pre-landing and post-landing muscle activation levels. Simultaneous recording of the jumping motion and ground reaction force was performed by a high-speed video camera (100 frames·s^–1), and a force platform was employed to allow joint moment analysis. Joint stiffness was calculated by a linear regression of the knee joint moment/angle relationship. Elasticity of the knee extensor muscle during DJ was estimated by means of a four-element muscle model consisting of a parallel elastic component, a series elastic component (SEC), a viscous damper, and a contractile element. DJ performance correlated positively with the positive peak power of the knee joint (P<0.01) and with the moment of the knee joint at the end of stretch (P<0.01). However, there was no significant relationship between DJ performance and the positive peak power of the ankle joint. The knee joint moment at the end of stretch correlated with the SEC stiffness during the transmission phase from the end of the initial impact to the onset of the concentric action (P<0.01) and with the maximum rate of isometric force development of the knee extensors (P<0.01). Multiple regression analysis showed that the SEC stiffness during the transmission phase of the knee joint can be explained by a combination of the pre-activity of the VL muscle and the knee joint angular velocity at touchdown (F=5.76, P<0.05). These results seem to emphasize the functional significance of the pre-programmed activity for controlling the subsequent stiffness regulation and then contributing to the performance in DJ. Thus, it can be suggested that the centrally pre-programmed activity and the associated elastic behavior of the SEC in the knee extensor muscle in conjunction with the muscle contractile property play a major role in regulating the performance in DJ. Electronic Publication     

Combined effect of elastic energy and myoelectrical potentiation during stretch-shortening cycle exercise

In addition to the utilization of muscle's elastic energy enhancement of performance in exercise involving stretch-shortening cycle might be also due to simultaneous increase of myoelectrical activity. This hypothesis was tested by examining three athletes during jumping exercise on force-platform. Vertical jumps were performed with and without preliminary counter-movement, and the jumps were called counter-movement jump (CMJ) and squatting jump (SJ), respectively. In both conditions several jumps were performed also with extra loads on the shoulders (15–220% of b. wt.). Additional droppingjumps (DJ) were executed from different heights (20–100 cm). During jumping exercise myoelectrical activity of selected muscles from the quadriceps femoris was monitored with surface electrodes. The results obtained were similar to those reported in isolated muscle and as expected, the prestretch in CMJ shifted the force-velocity curve of concentric work to the right. In two cases enhancement of performance was attributed primarily to restitution of elastic energy because myoelectrical activity was similar to that observed in SJ. In one subject increased myoelectrical activity was observed during the concentric phase of CMJ. In DJ condition the EMG activity during eccentric phase was much higher than in SJ. Therefore the high performance in this condition was attributed to both elastic energy and reflex potentiation. In eccentric work of CMJ the average force decreased with the increase of stretching speed. This phenomenon was associated with a light increase of EMG activity. The observed results emphasize that both elastic energy and reflex potentiation may operate effectively during stretch-shortening cycle activity.     

The mechanical efficiency of locomotion in men and women with special emphasis on stretch-shortening cycle exercises

Summary The mechanical efficiency of the leg extensor musculature of men and women was examined with a special sledge ergometer. The subjects (ten males and ten females) performed (a) pure positive work, (b) pure negative work and (c) a combination of negative and positive work (strech-shortening cycle). The mechanical efficiency of pure positive work was on average 19.8±1.2% for female subjects and 17.4±1.2% for male subjects (t=4.12, P<0.001), although the work intensity was equal in both groups. The mechanical efficiency of pure negative work was slightly lower in women than in men (59.3±14.4% vs 75.6±29.3%). The mechanical efficiency of positive work (₊) in a stretch-shortening cycle exercise was 38.1±6.8% in men and 35.5±6.9% in women. The utilization of prestretch was better for female subjects at low prestretch levels, whereas males showed greater potentiation of elastic energy at higher prestretch levels. Regarding absolute W _el (work due to elasticity) values, male subjects showed greater (P<0.001) values than females (189±44 J vs 115±36 J, respectively). Fundamental differences in neuromuscular functions in men and women might cause the differences in the results obtained.     

Effects of stretch-shortening cycle training on mechanical properties and fibre type transition in the rat soleus muscle

The effects of exercise training on mechanical properties and fibre type transitions have been investigated in rat soleus muscle. The exercise was a repetition of stretch-shortening cycles. A method of dual controlled releases was applied to obtain tension/extension curves, which characterize the elastic behaviour of the series elastic component (SEC), and the force/velocity relationship characterizing the contractile elements. Other contractile measurements included: contraction time (t _c), half-relaxation time (t _1/2) and twitch/tetanus ratio (P _t/P _o). Additionally, the muscle fibre type composition was determined by a classical histochemical method. A 12-week period of training induced a significantly higher percentage of fast-twitch fibres and a lower percentage of slow-twitch fibres (P<0.01). This fibre adaptation led to a significant (P<0.01) decrease in t _c and an increase in maximum shortening velocity (V _max). An increase in compliance of the SEC was also observed. This elastic adaptation is interpreted in terms of modification of the active components of the SEC. All the histochemical and mechanical data presented in this study show that rat soleus muscles trained by stretch-shortening cycles acquired faster characteristics. Thus the results confirm that a slow-twitch to a fast-twitch fibre transition is obtainable in mature rats.     

Changes in hardness of the human elbow flexor muscles after eccentric exercise

The purpose of this study was to investigate changes in muscle hardness after eccentric exercise of the elbow flexors muscles that produce muscle shortening and swelling. To assess muscle hardness, a pressure method was used in which the force required to deform the tissue (skin, subcutaneous tissue, muscle) was recorded. Eleven healthy male students performed 24 maximal eccentric actions of the elbow flexor muscles with their non-dominant arms. Muscle hardness, maximal isometric force (MIF), muscle soreness, plasma creatine kinase (CK) activity, relaxed elbow joint angle (RANG), upper-arm circumference (CIR) and B-mode ultrasound transverse images were measured before, immediately after, and 1–5 days after exercise. A long-lasting decrease in MIF, muscle swelling shown by increases in CIR and muscle thickness, large increases in plasma CK activity, and development of muscle soreness indicated that damage occurred to the elbow flexor muscles. The RANG had decreased by approximately 20° at 1–3 days after exercise and showed a gradual recovery thereafter. The CIR increased gradually after exercise and peaked on day 5 post-exercise, the mean amount of increase in CIR being 18 mm. Muscle hardness measured at the relaxed elbow position did not change until 3 days after exercise, but increased significantly (P < 0.01) on days 4 and 5 post-exercise. On the other hand, muscle hardness measured when forcibly extending the shortened elbow joint increased significantly (P < 0.01) with time and peaked at 3 days after exercise. Muscle hardness assessed by the pressure method seems to reflect changes in muscle stiffness and swelling. Accepted: 15 April 2000     

Ventilatory and circulatory responses at the onset of exercise after eccentric exercise

The purpose of this study was to clarify whether delayed onset muscle soreness (DOMS) and muscle damage after eccentric exercise (ECC) could affect the ventilatory and circulatory responses at the onset of exercise, and whether those effects would continue after the disappearance of DOMS. Ten males participated in this study. We measured ventilatory and circulatory responses at the onset of exercise, for the first 20 s, during knee extension–relaxation voluntary exercise (VOL) and passive movement (PAS), which was achieved by the experimenter alternatively pulling ropes connected to the subjects’ ankles for the same period and frequency as during VOL. VOL and PAS were performed before, 2 days after, and 7 days after ECC. The following results were found: (1) the gain of minute ventilation at the onset of VOL at 2 days after ECC was significantly larger than that of before ECC; (2) the gain of minute ventilation at 7 days after ECC during both VOL and PAS was also enhanced significantly as compared to that of before ECC; and (3) heart rate and blood pressure responses were unchanged throughout the experimental period. In conclusion, ventilatory response at the onset of exercise is augmented during DOMS and EIMD after ECC and the enhanced ventilatory response continued after the disappearance of DOMS. It is suggested that enhanced ventilatory response during exercise after ECC is attributed to at least the changes in neural factors and that the mechanisms inducing these augmented ventilatory responses should be different during the period after ECC.     

The effect of prolonged submaximal exercise on gas exchange kinetics and ventilation during heavy exercise in humans

This study compared ventilation, gas exchange (oxygen uptake, V̇O₂) and the surface electromyogram (EMG) activity of four major lower limb muscles during heavy exercise before (Pre-Ex) and after (Post-Ex) a sustained 90-min cycling exercise at 60% V̇O_2peak. The 90-min exercise was incorporated under the hypothesis that sustained exercise would alter substrate availability in the second exercise bout causing differences in fibre recruitment patterns, gas exchange and ventilation. Nine trained male subjects [V̇O_2peak=60.2 (1.7) ml·kg⁻¹·min⁻¹] completed two identical 6-min bouts of cycling performed at high intensity [~90% V̇O_2peak; 307 (6) W, mean (SE)]. Ventilation and gas exchange were measured breath-by-breath and the EMG was recorded during the last 12 s of each minute of the two 6-min bouts. EMG signals were analysed to determine integrated EMG (iEMG) and mean power frequency (MPF). V̇O₂ at min 3 and min 6 in Post-Ex were significantly higher (i.e., +201 and 141 ml·min⁻¹, respectively, P<0.05) than in Pre-Ex but there was a ~25% decrease of the slow component, taken as the difference between min 6 and min 3 [187 (27) vs 249 (35) ml·min⁻¹, respectively, P<0.05]. The greater whole-body V̇O₂ after 3 min of exercise in Post-Ex was not accompanied by clear alterations in the iEMG and MPF of the examined leg muscles. Ventilation and heart rate were elevated (~12–16 l·min⁻¹ and ~10 beats·min⁻¹, respectively, P<0.05) as were the ratios V̇ _E/O₂ and V̇ _E/V̇CO₂ in the Post-Ex tests. It was concluded that the V̇O₂ and ventilation responses to high-intensity exercise can be altered following prolonged moderate intensity exercise in terms of increased amplitude without associated major changes in either iEMG or MPF values among conditions.     

Carbohydrate supplementation and alterations in neutrophils, and plasma cortisol and myoglobin concentration after intense exercise

The present study examined the effect of carbohydrate supplementation on changes in neutrophil counts, and the plasma concentrations of cortisol and myoglobin after intense exercise. Eight well-trained male runners ran on a treadmill for 1 h at 85% maximal oxygen uptake on two separate occasions. In a double-blind cross-over design, subjects consumed either 750 ml of a 10% carbohydrate (CHO) drink or a placebo drink on each occasion. The order of the trials was counter-balanced. Blood was drawn immediately before and after exercise, and 1 h after exercise. Immediately after exercise, neutrophil counts (CHO, 49%; placebo, 65%; P<0.05), plasma concentrations of glucose (CHO, 43%; P<0.05), lactate (CHO, 130%; placebo, 130%; P<0.01), cortisol (CHO, 100%; placebo, 161%; P<0.01), myoglobin (CHO, 194%; placebo, 342%; P<0.01) all increased significantly. One hour post-exercise, plasma myoglobin concentration (CHO, 331%; placebo, 482%; P<0.01) and neutrophil count (CHO, 151%; placebo, 230% P<0.01) both increased further above baseline. CHO significantly attenuated plasma myoglobin concentration and the neutrophil count after exercise (P<0.01), but did not affect plasma cortisol concentration. The effects of CHO on plasma myoglobin concentration may be due to alterations in cytokine synthesis, insulin responses or myoglobin clearance rates from the bloodstream during exercise. Plasma cortisol responses to CHO during exercise may depend on the intensity of exercise, or the amount of CHO consumed. Lastly, cortisol appears to play a minor role in the mobilisation of neutrophils after intense exercise.     

Muscular function,metabolism and electrolyte shifts during prolonged repetitive exercise in humans

Marked functional changes occur in human skeletal muscle during prolonged repetitive exercise. The maximum voluntary contraction force (MVC) falls gradually and may reach 50% of control within 30–60 min. The twitch tension declines faster and to a larger extent. During repetitive submaximal isometric contractions, the rate of relaxation increases progressively, in parallel with an increased energy cost of contraction. These functional changes are all slowly reversed in the post-exercise period, as indicated by only minor changes over the first 30 min of recovery. Minor changes in substrates and metabolites, together with the slow rate of recovery, indicate that the alterations in contractile properties and energetics are independent of these metabolic factors. Alternative explanations may be related to electrolyte shifts over the sarcolemma or between cellular compartments. The total loss of K⁺ is small, and could not be detected by analysis of muscle biopsies. Only a slight initial rise in muscle content of calcium was found. The available data indicate that the increased energy cost of contraction is not connected to mitochondrial dysfunction, which might be caused by calcium accumulation. Rather, it seems that the ratio of ATP utilization to force is increased, and this could possibly be connected to the faster relaxation rate. Considering the low excitation rates during submaximal voluntary contractions, each motor unit generates an oscillating force closely associated with Ca²⁺ fluctuations between SR and cytosol. Increased relaxation rate might be caused by faster reuptake of Ca²⁺ into the SR, and this could contribute to the faster ATP turnover.     

Alterations in the expression and activity of creatine kinase-M and mitochondrial creatine kinase subunits in skeletal muscle following prolonged intense exercise in rats

Creatine kinase (CK) isoenzymes are important structural and energy metabolism components in skeletal muscle. In this study, CK isoenzyme alterations were examined in male rats, with an 8% body mass weight attached to their tail. The rats were either forced to swim for 5 h (5S, n = 51), or were pre-trained for 8 days and then forced to swim for 5 h (T5S, n = 48). Rats were sacrificed either immediately (0 h PS), 3 h (3 h PS), or 48 h post-swimming (48 h PS). Serum CK was increased significantly (P < 0.01) 6.2- and 2.0-fold at 0 h PS following the 5S and T5S protocols, respectively. However, training (T5S protocol) significantly (P < 0.01) decreased CK release. Soleus and white gastrocnemius (WG) CK activity was significantly decreased following the 5S protocol (P < 0.05), but not following the T5S protocol. The CK-M activity of the soleus muscle was significantly (P < 0.05) decreased at 0 h PS following both the 5S and T5S protocols, and returned to control values at 3 h PS. The CK-M activity of the WG was significantly (P < 0.05) decreased at 0 h PS following the 5S protocol. Sarcomeric mitochondrial CK (sCK-Mit) was decreased significantly (P < 0.01) at 0 h PS (20%), 3 h PS (14%), 24 h PS (22%), and 48 h PS (15%) following the 5S protocol. However, sCK-Mit was decreased significantly (P < 0.01) only at 0 h PS (7%) following the T5S. The results of this study demonstrate that prolonged intense exercise causes a loss of skeletal muscle CK-M and sCK-Mit activity and that training prior to the prolonged intense exercise attenuates the exercise-induced CK-M and sCK-Mit loss in both red and white skeletal muscles. Accepted: 18 July 1999     

Temporal summation of pain from skin,muscle and joint following nociceptive ultrasonic stimulation in humans

This study investigated the phenomenon of temporal summation in response to repetitive focused ultrasound stimulation of skin, muscle and joint in human volunteers. Stimulation was carried out using a custom-designed, focused ultrasonic stimulator with a resonant frequency of 1.66 MHz. A series of stand-off attachments were used to ensure that the focal region of the ultrasound beam projected either cutaneously, within the distal interphalangeal joint of the index finger, or within the first dorsal interosseous muscle. Stimulation was carried out using single pulses and trains of five pulses of different pulse durations (25 ms, 50 ms, 75 ms, 100 ms), and using single pulses and trains of five pulses (50 ms duration) at different frequencies (0.5 Hz, 1 Hz, 2 Hz, 3 Hz, 4 Hz, 5 Hz). Tactile perception thresholds, pain thresholds and summation pain thresholds were recorded. Temporal summation of pain could be elicited by stimulation of both skin, joint and muscle, although the influence of temporal summation appeared to be more pronounced for muscle stimulation. Muscle stimulation also required greater ultrasound intensity compared with joint and skin stimulation. Temporal summation could not be elicited by tactile, low-intensity stimulation. Focused ultrasound is a potent, noninvasive technique with which to investigate temporal summation from somatic structures. A number of factors may account for the higher intensities required to elicit pain in muscle and the increased rate of temporal summation. It is clear, however, that if temporal summation is more pronounced in muscle than other tissues then this may be an important factor contributing to pain in musculoskeletal syndromes.