Age-related changes in power output during repetitive contractions of rat medial gastrocnemius muscle

Changes in isometric force, power output and relaxation rate have been measured during repetitive tetanic contractions in 2 groups of rats of different ages. During the first 5 contractions there were no differences between a young and mature group. In contrast to isometric force production, which decreased about 3% per contraction, power output initially increased to 108% of the power output in the first contraction. A greater reduction in power output and relaxation rate after the 5th contraction indicated a greater reduction of the cross-bridge cycling rate in the younger rats. ATP, phosphocreatine and lactate concentrations after the last contraction were not different between the age-groups. In contrast IMP production, which has been suggested may play a regulatory role during fatigue was twice as high in the young rats. Judged by isometric force production there is no age-related difference in fatiguability. However, profound differences were observed in power output, which indicates that quantification of fatigue as a loss of isometric force may be seriously misleading when considering the functional status of the muscle for normal dynamic contractions.     

Age-related changes in force and efficiency in rat skeletal muscle

We investigated the effect of age on (the reduction of) work output, efficiency and muscle fibre type composition. Rat medial gastrocnemius muscles of three age-groups performed a series of 15 repeated contractions within 6 s (blood flow was arrested). Stimulation and shortening velocities were chosen as optimal for each group, while all muscles shortened over the same relative fibre lengths. The fibre type composition showed a higher proportion of the oxidative type IIBd fibres in the middle-aged group [5 months old; 39.8 ± 6.8 vs. 23.6 ± 4.2% of the fibre area in the young rats (1.3 months old)] in contrast to the type IIBm fibres (52.9 vs. 67.9%, respectively), while the old group (22 months old) was not different from the middle-aged group. Work output in the last contraction (relative to the first contraction) was not different between the age-groups (53.1 ± 18.1; 48.± 6.5 and 61.1± 6.2%, respectively). High-energy phosphate utilization was not different between the groups (150.6 ± 11.2; 154.6 ± 15.6 and 157.2 ± 7.0 μmol g^-1 dry wt, respectively). However, the efficiency was 30% lower in the muscles of the youngest group, which corresponds with a lower specific power and specific tension. Since the change in fibre type composition is unlikely to be the cause of the low efficiency in the young animals, the causes remain unclear, but may be related to the rapid growth of the young rats in our study.     

Anaerobic power output of young obese men: Comparison with non-obese men and the role of excess fat

Summary Anaerobic power output was measured by the staircase climb test in 14 obese, 16 lean, and 21 ordinary men aged from 18–22 years. Fat storage rate (%fat) was estimated by densitometry. The obese group ranked highest with an average power output of 1,012 W. This value was significantly higher than those of the other two groups, 890 W for lean subjects and 855 W for ordinary subjects. The power output per kilogram of lean body mass of the obese group was the highest also. However, the vertical velocity was the lowest although the difference among the three average values was not statistically significant. To investigate the effect of excess fat, eight non-obese subjects engaged in an added-weight experiment. The value obtained was almost the same as for the obese group. The added weights made the vertical velocity decrease but the power output increase. Consequently, it was obvious that the excess fat of an obese man played a role only as an inert mass in the power output measurement. A significantly higher power output of the obese group might be due to more excess fat, and obesity itself was an advantage in the staircase climb test.     

The effect of repeated muscle biopsy sampling on ATP and glycogen resynthesis following exercise in man

The present study investigated the effect of repeated biopsy sampling on muscle adenosine 5-triphosphate (ATP) and glycogen resynthesis following prolonged submaximal exercise. In one group of subjects (Ia, n = 7), biopsy specimens were obtained from the vastus lateralis immediately and 48 h after exhaustive one-legged cycling from both the non-exercised (control) and exercised legs. Additional samples were obtained from the exercised leg at 3, 10 and 24 h post-exercise. In a second group of subjects (Ib, n = 6), biopsy specimens were obtained immediately after exercise from both the control and exercised legs and at 48 h post-exercise from the exercised leg. All muscle biopsies were separated by a distance of 2.5 cm. In group Ia, ATP in the exercised leg was still lower after 48 h of recovery compared with the control leg (P < 0.05), but complete restoration had occurred in group Ib (P > 0.05). Glycogen super compensation was not observed in group Ia. However, at the end of recovery, in group Ib glycogen in the exercised leg was 42% greater than in the control leg (tP < 0.01) . Thus, following exhaustive dynamic exercise, repeated muscle biopsy sampling impaired ATP and glycogen resynthesis for several days, which may have been a result of the distance separating each biopsy site. The inhibition of ATP resynthesis appeared to be associated mainly with type II muscle fibres. The finding that, in contrast to muscle glycogen, ATP did not return to the basal level during the 48 h of recovery, suggests that the measurement of ATP may be a more sensitive measure of muscle damage than that of glycogen.     

Heat production and chemical change during isometric contraction of rat soleus muscle.

1. Methods are described whereby the soleus muscle of the rat may be used for the investigation of initial processes in the absence of oxidative recovery. 2. The anaerobic conditions employed had no effect on the concentration of phosphocreatine in resting muscle or the mechanical response during contraction. 3. Muscles were stimulated tetanically for 10 s at 17-18 degrees C. Measurements were made of the heat production and metabolic changes that occurred during a 13 s period following the first stimulus. 4. There was no detectable change in the concentration of ATP. Neither was there detectable activity of adenylate kinase or adenylate deaminase. The changes in the concentration of glycolytic intermediaries were undetectable or very small. 5. The change in the concentration of phosphocreatine was large and amounted to -127 +/- 11-4 mumol/mmol Ct (mean and S.E. of the mean, negative sign indicates break-down, Ct = free creatine + phosphocreatine) which is equivalent to about -2-13 mumol/g wet weight of muscle. The heat production was 6549 +/- 408 mJ/mmol Ct (mean and S.E. of mean) which is equivalent to about 110 mJ/g. 6. About 30% of the observed energy output is unaccounted for by measured metabolic changes. 7. The ratio of heat production (corrected for small amounts of glycolytic activity) to phosphocreatine hydrolysis was -49-7 +/- 5-6 kJ/mol (mean and S.E. of mean), in agreement with previous results using comparable contractions of frog muscle, but different from the enthalpy change associated with phosphocreatine hydrolysis under in vivo conditions (-34 kJ/mol). 8. The results support the notion that the discrepancy between energy output and metabolism is an indication of an unidentified process of substantial energetic significance that is common to a number of species.     

Anaerobic ATP provision,glycogenolysis and glycolysis in rat slow-twitch muscle during tetanic contractions

Rat soleus muscles were tetanically stimulated in situ with an occluded circulation to examine anaerobic adenosine triphosphate (ATP) provision and the regulation of glycolytic ATP production. Soleus muscles were stimulated for 30–60 s at 1 Hz with 100–200 ms trains (40–80 Hz). Muscles were sampled pre- and post-stimulation for measurements of pH, high energy phosphates and glycolytic intermediates. Total ATP provision by the slow oxidative fibres was 65–121 mol/g dry muscle and 27–35% of the amount produced by fast glycolytic fibres. Contributions to total ATP provision in the initial 30 contractions were: phosphocreatine, 71%; glycolysis, 28%; and endogenous ATP, 1%. Following 60 contractions the contributions were 45–54%, 44–51% and 2–4%, respectively. During the initial 30 contractions, glycogenolysis (phosphorylase activity) and glycolysis [phosphofructokinase (PFK) activity] were similar as glucose-6-phosphate (G-6-P) and fructose-6-phosphate (F-6-P) did not accumulate. Small accumulations of PFK deinhibitors inorganic phosphate, adenosine diphosphate, adenosine monophosphate and fructose-1,6-diphosphate appeared to account for the PFK activity. In the final 30 contractions, phosphorylase activity increased above PFK as G-6-P and F-6-P accumulated. PFK activity and glycolytic ATP production also increased despite increasing hydrogen ion concentration [H⁺]. During intense tetanic stimulation of soleus muscle, glycolytic ATP production is initially limited by a low glycogen phosphorylase activity. The activity of PFK increases during in situ contraction through the accumulation of deinhibitors, despite increasing [H⁺].     

Heat production and chemical changes during isometric contractions of the human quadriceps muscle.

1. Development of a new thermal probe and use in conjunction with chemical analysis of needle biopsy samples, has made possible a thermodynamic study of the energetics of muscular contraction in the human quadriceps. 2. The observed rate of muscle temperature rise was proportional to the force of the contraction. During maximal contractions the rate of heat production was 54 +/-8-5 W/kg wet muscle (mean +/- s.d.). 3. The observed rates of muscle temperature rise agreed well with the rates calculated from the measured metabolite changes when standard values for the enthalpy changes of the reactions involved were used. 4. During prolonged stimulation of the quadriceps at 15/sec via the femoral nerve, the rate of heat production per unit force fell to nearly half the initial value. It is estimated that this represented a two- to fourfold increased in economy of ATP turn-over required to maintain a given force. 5. Relaxation becomes progressively slower during prolonged contractions and it is suggested that the slowing of relaxation and the increased economy of force maintenance may both be due to an increased cross-bridge cycle time in the fatigued muscle.     

The time course of the last contractions during incompletely fused tetani of motor units in rat skeletal muscle

The time course of the contraction and the relaxation of individual contractions during incompletely fused tetani of motor units were analyzed. Investigations were performed on fast fatigable (FF), fast resistant (FR) and slow (S) motor units of the rat medial gastrocnemius muscle. Stimulation of a motor unit with a series of nine trains of stimuli at a frequency from 10 to 150 Hz was used and tetani fused to a variable degree were recorded. For fast motor units the procedure was repeated twice and observations were made on potentiated tetani in the second series of stimulation. For each tetanus, the amplitude of the tension increase, the peak amplitude of the contraction, the contraction time and the half-relaxation time were measured in the last contraction of the tension recording. It was observed in all three types of motor units that the last contraction was prolonged in parallel with the increase of fusion of a tetanus. In this contraction, the contraction time slightly decreased whereas the half-relaxation time strongly prolonged. The prolongation of the half-relaxation time was the strongest in tetani of slow units. Moreover, for the last contraction in a tetanus, the rate of changes in tension were studied. The rate of increase in tension during the contraction decreased in parallel with the increase of fusion of a tetanus, whereas the maximal rate of the tension decrease during the relaxation was found in tetani with fusion indices of 0.79, 0.98 and 0.95 for FF, FR and S motor units, respectively. Changes in the time course of contractions in tetani fused to a variable degree can shed light on processes of summation of contractions in unfused tetani at the level of individual motor units.     

Anaerobic threshold,skeletal muscle enzymes and fiber composition in young female cross-country skiers

Anaerobic threshold (AT) and maximum oxygen uptake (max V?_o2) were determined in 15 young female cross-country skiers, aged 15–20 years, during incremental bicycle ergo-meter exercise. Succinate dehydrogenase (SDH), malate dehydrogenase (MDH), citrate synthase (CS) and lactate dehydrogenase (LDH) were analyzed biochemically and percentage of slow twitch fibres (%ST fibres, myosin adenosine triphosphatase staining) histochemically in muscle samples obtained from m. vastus lateralis. Max V?_o2 correlated significantly with anaerobic threshold in ml×kg^-1×min^-1 (mlAT) but when AT was expressed in percent of max V?o2 (%AT) the correlation was insignificant. Significant correlations were found between %AT and SDH (r=0.63) and between mlAT and CS (r=0.58). Max V?_o2 showed no significant correlations with the enzymes studied or %ST fibres. The results of the study seem to support the hypothesis that anaerobic threshold is related to oxidative capacity of muscle.     

Mechanical and electrical correlates of isometric muscle fatigue in skeletal muscle in the cat

The amplitude and duration of motor unit action potentials, the rise time, peak tension, and half relaxation time of an isometric twitch, and forcevelocity relationship, and tetanic tension were measured at the beginning (fresh muscle) and at the end of a fatiguing isometric contraction at a tension of either 40 or 70% of the initial strength in the soleus (a slow twitch muscle) and the medial gastrocnemius (a fast twitch muscle) of the cat. These same parameters were also measured at set intervals following these contractions to assess their rate of recovery to pre-exercise values. At The end of a fatiguing contraction examined,V _mx, twitch tension and tetanic tension, were all reduced while there was a prolonged twitch duration and duration of the motor unit action potential for both types of muscle. The height of the motor unit action potential was only marginally effected by muscle fatigue. Following the fatiguing contraction, the endurance required several hours to recover in the medial gastrocnemius muscle but recovered fully within 15 min after either tension in the soleus muscle. Tetanic tension and twitch tension both required less than 10 min for full recovery in the medial gastrocnemius muscle but recovered fully to the pre-exercise values within 3 min following fatiguing isometric contractions in the soleus muscles.V _mx, and the height and duration of the motor unit action potential both recovered within 1 min following the end of the exercise.     

Changes in velocity of shortening,power output and relaxation rate during fatigue of rat medial gastrocnemius muscle

The force-velocity characteristics of rat medial gastrocnemius muscle have been determined by measuring the force sustained during constant velocity releases of the muscle stimulated in situ at an ambient temperature of 26°C. The velocity of unloaded shortening was determined using the slack test and rate of relaxation from the half time of force loss at the end of stimulation. Measurements were first made on fresh muscles using short contractions and then during a series which consisted of a 15 s contraction (fatigued muscle), followed by 15 min recovery and a 1 s contraction (recovered muscle). After a 5 min recovery period the sequence was repeated. Comparison was made between the fatigued and recovered state in each preparation in order to allow for any change in the preparation during the course of the experiment. After 15 s contraction the fatigued muscles showed a marked reduction in all parameters measured. In fatigued muscles the isometric force fell to 48±15% (mean±SD) and there was a decrease in maximum velocity of shortening to 66%. These changes in the force-velocity relationship were accompanied by slowing of relaxation so that the half time of relaxation nearly doubled. The consequence of these changes was that the maximum power output was reduced by a much greater extent that was the isometric force (75% vs. 52%). It is suggested that the changes in force-velocity characteristics reflect a reduction in cross-bridge cycling in fatigued muscle.     

Age- and training-related changes in the collagen metabolism of rat skeletal muscle

Summary The effects of ageing and life-long endurance training on the collagen metabolism of skeletal muscle were evaluated in a longitudinal study. Wistar rats performed treadmill running 5 days a week for 2 years. The activities of collagen biosynthesis enzymes, prolyl-4-hydroxylase and galactosylhydroxylysyl glucosyltransferase, were highest in the muscles of the youngest animals, decreased up to the age of 2 months and from then on remained virtually unchanged. The enzyme activity in young animals was higher in the slow collagenous soleus muscle than in the rectus femoris muscle. The enzyme activity in the soleus muscle was higher for older trained rats than older untrained rats. The relative proportion of type I collagen increased and that of type III collagen decreased with age, suggesting a more marked contribution by type I collagen to the agerelated accumulation of total muscular collagen. The results show that collagen biosynthesis decreases with maturation and that life-long endurance training maintains a higher level of biosynthesis in slow muscles.     

Function dependent changes in the subcellular distribution of high energy phosphates in fast and slow rat skeletal muscles

Function dependent changes in the subcellular distribution of ATP, ADP, creatine phosphate (CrP) and creatine (Cr) in rat fast-twitch gastrocnemius and slow-twitch soleus muscles were studied by fractionation of freeze-clamped and freeze-dried tissue in non-aqueous solvents.During 5 min of isotonic contraction of gastrocnemius muscles the mitochondrial content of total creatine [(CrP+Cr)] decreases by 9.5 nmol/mg total protein whereas there is an increase in extramitochondrial total creatine by 12.3 nmol/mg total protein, indicating a net transfer of 10 nmol total creatine/mg total protein/5 min across the mitochondrial inner membrane.During short-term stimulation (6 s) of gastrocnemius muscles the socalled additionally-bound ADP correlates not only with force (Hebisch et al. 1984) but also with filament overlap. This confirms the previous suggestion that additionally-bound ADP represents actomyosin-ADP-complexes.Following long-term stimulation (10 s), the rate of decay of force is at least two orders of magnitude faster than that of additionally bound ADP. This indicates a decrease of actomyosin-ADP complexes due to formation of myosin-ADP complexes.Short-term stimulation (6 s) of slow-twitch soleus muscles does not lead to any force-dependent change in the content of additionally-bound ADP, similar to the finding in long-term contracting gastrocnemius muscles. Denervation of soleus muscles leads to a decrease in additionally-bound ADP to values comparable to those found in resting fast-twitch gastrocnemius muscles.     

Metabolic cost of lengthening, isometric and shortening contractions in maximally stimulated rat skeletal muscle

AIM: The present study investigated the energy cost of lengthening, isometric and shortening contractions in rat muscle (n = 19). METHODS: With electrical stimulation the rat medial gastrocnemius muscle was maximally stimulated to perform 10 lengthening, isometric and shortening contractions (velocity 25 mm s(-1)) under experimental conditions (e.g. temperature, movement velocity) that resemble conditions in human movement. RESULTS: Mean +/- SD force-time-integral of the first contraction was significantly different between the three protocols, 2.4 +/- 0.2, 1.7 +/- 0.2 and 1.0 +/- 0.2 N s, respectively (P < 0.05). High-energy phosphate consumption was not significantly different between the three modes of exercise but a trend could be observed from lengthening (7.7 +/- 2.7 micromol approximately P muscle(-1)) to isometric (8.9 +/- 2.2 micromol approximately P muscle(-1)) to shortening contractions (10.4 +/- 1.6 micromol approximately P muscle(-1)). The ratio of high-energy phosphate consumption to force-time-integral was significantly lower for lengthening [0.3 +/- 0.1 micromol approximately P (N s)(-1)] and isometric [0.6 +/- 0.2 micromol approximately P (N s)(-1)] contractions compared with shortening [1.2 +/- 0.2 micromol approximately P (N s)(-1)] contractions (P < 0.05). CONCLUSION: The present results of maximally stimulated muscles are comparable with data in the literature for voluntary human exercise showing that the energy cost of force production during lengthening exercise is approximately 30% of that in shortening exercise. The present study suggests that this finding in humans probably does reflect intrinsic muscle properties rather than effects of differential recruitment and/or coactivation.     

The mechanical response of active human muscle during and after stretch

Summary Five subjects contracted forearm supinator muscles which were stretched after development of maximal isometric torque. The ratio of torque at the end of stretch over isometric torque at that position was calculated as excess torque. Excess torque increased with stretch velocity and decreased with stretch amplitude, and it was not dependent upon final muscle length. The rate of decay of torque following stretch could not be shown to depend upon stretch variables.The absence of significant changes in myoelectric activity suggested that with high initial forces, reflex activity did not account for the observed changes. Time-constants of decay (0.15 s to 1.8 s) were much greater than time-constants of rise (approx. 0.07 s) of isometric torque at the same muscle length. This indicates that interaction of series elastic and contractile elements is not the sole cause of prolonged torque following stretch. It is concluded that stretch temporarily enhances the intrinsic contractile properties of a group of human muscles in a manner similar to, but quantitatively different from that seen in isolated muscle preparations.     

Stoichiometry of phosphocreatine and inorganic phosphate changes in rat skeletal muscle

The apparent T1's of phosphate metabolites were measured during and after series of twitch contractions in gastrocnemius muscles of pentobarbital-anesthetized rats by steady-state progressive saturation using spatially-selective composite pulses. There was no significant change in apparent T1's of inorganic phosphate (Pi), phosphocreatine (PCr), or the three phosphates of ATP. There was a 5-10% decrease in the sum of Pi and PCr integrals, and in the sum of the gamma- and beta- phosphates of ATP during stimulation, but no significant change in the ratio (Pi + PCr)/(gamma-ATP + beta-ATP). The results indicate that there is no selective decrease in NMR observable Pi during or after a series of muscle contractions.     

Effects of isokinetic training of the knee extensors on isometric strength and peak power output during cycling

Summary Isokinetic training of right and left quadriceps femoris was undertaken three times per week for 16 weeks. One group of subjects (n=13) trained at an angular velocity of 4.19 rad·s^–1 and a second group (n=10) at 1.05 rad·s^–1. A control group (n=10) performed no training. Maximal voluntary contraction (MVC) of the quadriceps, and peak pedal velocity (_p,peak) and peak power output (W _peak) during all-out cycling (against loads equivalent to 9, 10, 11, 12, 13 and 14% MVC) were assessed before and after training. The two training groups did not differ significantly from each other in their training response to any of the performance variables (P>0.05). No significant difference in MVC was observed for any group after the 16-week period (P=0.167). The post-training increases in averageW _peak (7%) and _p,peak (6%) during the cycle tests were each significantly different from the control group response (P=0.018 andP=0.008, respectively). It is concluded that 16 weeks of isokinetic strength training of the knee extensors is able to significantly improve _p,peak andW _peak during sprint cycling, an activity which demands considerable involvement of the trained muscle group but with its own distinct pattern of coordination.     

Impact of post-synaptic block of neuromuscular transmission, muscle unloading and mechanical ventilation on skeletal muscle protein and mRNA expression

To analyse mechanisms of muscle wasting in intensive care unit patients, we developed an experimental model where rats were pharmacologically paralysed by post-synaptic block of neuromuscular transmission (NMB) and mechanically ventilated for 9±2 days. Specific interest was focused on the effects on protein and mRNA expression of sarcomeric proteins, i.e., myosin heavy chain (MyHC), actin, myosin-binding protein C (MyBP-C) and myosin-binding protein H (MyBP-H) in fast- and slow-twitch limb, respiratory and masticatory muscles. Muscle-specific differences were observed in response to NMB at both the protein and mRNA levels. At the protein level, a decreased MyHC-to-actin ratio was observed in all muscles excluding the diaphragm, whereas at the mRNA level a decreased expression of the dominating MyHC isoform(s) was observed in the hind limb and intercostal muscles, but not in the diaphragm and masseter muscles. MyBP-C mRNA expression was decreased in the limb muscles, but it otherwise remained unaffected. MyBP-H conversely increased in all muscles. Furthermore, we found myofibrillar protein and mRNA expression to be affected differently when comparing NMB animals with peripherally denervated (DEN) ambulatory rats. We report that NMB has both a larger and different impact on muscle, at the protein and mRNA levels, than DEN has.     

Influence of muscle dimensions on economy of isometric exercise in rat medial gastrocnemius muscles in situ

Summary The effect of muscle dimensions on economy (force-time integral divided by the amount of energy utilized) was investigated in male rats (body mass range 95–490 g), anaesthetized with pentobarbital. The medial gastrocnemius muscle in situ performed 6 maximal isometric contractions of 350 ms duration (1 · s^–1) at twitch optimum length at 35°C. The areas under the 6 time-force curves were added to obtain force-time integral of the experiment. Differences of concentrations of ATP, phosphocreatine and lactate between experimental and contralateral (resting) muscles were used to calculate high-energy phosphate consumption due to stimulation. Muscle mass and cross-sectional area increased (approximately +400% and +300%, respectively) over the rat body mass range studied. Muscle length and length of the most distal fibre bundle increased by approximately 17 mm and 4 mm, respectively. Force-time integral (N · s) increased proportional to cross-sectional area whereas high-energy phosphate consumption (moles) increased proportional to muscle mass. The relative fraction of the total energy consumption utilized for force-independent processes was independent of rat body mass. The economy of the actomyosin system was unaffected during growth, whereas economy of the whole muscle decreased during growth by approximately 30% (p<0.001). The effect of muscle dimensions on economy is discussed with respect to human endurance capacity measured by voluntary isometric contractions.