This research carried out in Amsterdam under the direction of Professor Anthony J Sargeant demonstrated how within the same anatomical muscle there can be quiet different physiological properties in different areas of the same muscle. This work was part of the PhD research of Jo de Ruiter supervised by Professor Tony Sargeant and Arnold de Haan.
The most proximal and distal motor nerve branches in the rat medial gastrocnemius innervate discrete muscle compartments dominated by fast-twitch oxidative and fast-twitch glycolytic fibers, respectively. The functional consequences of the difference in oxidative capacity between these compartments were investigated. Wistar rats were anesthetized with pentobarbital sodium (90 mg/kg ip). Changes in force of both compartments during 21 isometric contractions (train duration 200 ms, stimulation frequency 120 Hz, 3 s between contractions) were studied in situ with and without blood flow. Without blood flow, force and phosphocreatine declined to a greater extent in the proximal than the distal compartment compared with the run with intact flow. After the protocol without blood flow, when flow was restored, the time constants for force recovery (which were closely associated to the recovery of phosphocreatine) were 37 +/- 7 (SD) (proximal compartment) and 148 +/- 20 s (distal compartment). It was concluded that the proximal compartment had a four times higher oxidative capacity and, therefore, a superior ability for repeated force production.
The ability of generating muscle power power is important whether you are an Olympic athlete, a ballet dancer, or an elderly person wanting to climb the stairs to go to bed. In this comprehensive review of his research Anthony Sargeant points out the importance of different types of muscle fibres that make up the human skeletal muscles that produce power in legs and arms. Tony also points out that in research seeking to measure human muscle power it is essential to measure or control the speed at which the power is generated (this is because power is the product of work and velocity).
Structural and functional determinants of human muscle power
by Anthony J Sargeant
Measurements of human power need to be interpreted in relation to the movement frequency, since that will determine the velocity of contraction of the active muscle and hence the power available according to the power-velocity relationship. Techniques are described which enable movement frequency to be kept constant during human exercise under different conditions. Combined with microdissection and analysis of muscle fibre fragments from needle biopsies obtained pre- and postexercise we have been able ‘to take the muscle apart’, having measured the power output, including the effect of fatigue, under conditions of constant movement frequency. We have shown that fatigue may be the consequence of a metabolic challenge to a relatively small population of fast fatigue-sensitive fibres, as indicated by [ATP] depletion to approximately 30% of resting values in those fibres expressing myosin heavy chain isoform IIX after just 10 s of maximal dynamic exercise. Since these same fibres will have a high maximal velocity of contraction, they also make a disproportionate contribution to power output in relation to their number, especially at faster movement rates. The microdissection technique can also be used to measure phosphocreatine concentration ([PCr]), which is an exquisitely sensitive indicator of muscle fibre activity; thus, in just seven brief maximal contractions [PCr] is depleted to levels < 50% of rest in all muscle fibre types. The technique has been applied to study exercise at different intensities, and to compare recruitment in lengthening, shortening and isometric contractions, thus yielding new information on patterns of recruitment, energy turnover and efficiency.
Anita Beelen presented this research as part of her PhD thesis supervised by Professor Anthony Sargeant. Uniquely the study used electrical stimulation superimposed upon on maximal voluntary activation in dynamic exercise.
1. Percutaneous electrical stimulation of the human quadriceps muscle has been used to assess the loss of central activation immediately after a bout of fatiguing exercise and during the recovery period.
2. Fatigue was induced in eight healthy males by a maximal effort lasting 25 s performed on an isokinetic cycle ergometer at a constant pedal frequency of 60 revolutions per minute. The cranks of the ergometer were driven by an electric motor. Before and after the sprint, subjects allowed their legs to be passively taken round by the motor. During the passive movement the knee extensors were stimulated (4 pulses; 100 Hz). Peak voluntary force (PVF) during the sprint and peak stimulated forces (PSF) before and in recovery were recorded via strain gauges in the pedals. Recovery of voluntary force was assessed in a series of separate experiments in which subjects performed a second maximal effort after recovery periods of different durations.
3. Peak stimulated forces were reduced to 69f8 + 9 3 % immediately after the maximal effort, (P< 0 05), but had returned to pre-exercise values after 3 min. The maximum rate of force development (MRFD) was also reduced following fatigue to 68f8 + 11 0% (P < 0’05) of control and was fully recovered after 2 min. PVF was reduced to 72-0 + 9 4% (P< 0 05) of the control value following the maximal effort. After 3 min voluntary force had fully recovered.
4. The effect of changing the duration of the fatiguing exercise (10, 25 and 45 s maximal effort) resulted in an increased degree of voluntary force loss as the duration of the maximal effort increased. This was associated with an increased reduction in PSF measured immediately after the exercise.
5. The close association between the changes in stimulated force and voluntary force suggests that the fatigue in this type of dynamic exercise may be due to changes in the muscle itself and not to failure of central drive.
This research was part of the PhD thesis of CJ (Jo) de Ruiter. It shows how within a single muscle there may be marked regional differences in physiological characteristics implying task dependent differences in recruitment patterns of motor units. The work was carried out under the direction of Anthony J Sargeant and Arnold de Haan.
Rat medial gastrocnemius (GM) muscle is a compartmentalized muscle. The functional properties and fibre type composition of the most proximal and most distal compartment were studied in in situ preparations. The proximal compartment contained predominantly fast twitch oxidative fibres. The distal compartment was mainly composed of fast twitch glycolytic fibres. With the use of two small electrodes placed around the primary nerve branches, both compartments could be separately stimulated within the same muscle. The length-force relationship was less broad and maximal twitch and tetanic forces were obtained at lower muscle lengths for the proximal compartment. The differences (mm) were 0.9 +/- 0.2 and 1.2 +/- 0.2 for maximal twitch and tetanic force (120 Hz) production, respectively (P < 0.001). The shortening velocity for maximal power production was lower (P < 0.001) for the proximal compartment (proximal: 57.1 +/- 2.7 mm s-1, distal: 73.1 +/- 3.0 mm s-1). During a standard fatigue test the fatiguability was significantly lower for the proximal compared with the distal fibres. Our findings suggest that the proximal compartment is likely to be activated in vivo during activities requiring relatively low power outputs for longer time periods. In contrast the distal compartment is probably recruited only during high power demanding short lasting activities. The presented model makes it possible to study fatigue related changes in power production of the ‘red’ and ‘white’ areas of the GM separately in a way that is probably meaningful with respect to in vivo function.
Neuromuscular function and fatigue have been studied using a wide variety of preparations. These range from sections of single fibers from which the cell membrane has been removed to whole muscles or groups of muscles acting about a joint in the intact animal. Each type of preparation has its merits and limitations. There is no ideal preparation; rather the question to be answered will determine the most appropriate model in each case and sometimes a combination of approaches will be needed. In particular, it is important to understand how the mechanical output of whole muscle can be sustained to meet the demands of a task and to take into account the organized variability of the constituent motor units.
The effect of growth on work output, energy consumption and efficiency during repetitive dynamic contractions was determined using extensor digitorum longus muscles of 40-, 60-, 120- and 700-day-old male Wistar rats. When work output of each contraction was normalized to the work output of the first contraction it was found that work output initially increased over the first 10-20 contractions by approximately 8% in each age group. Thereafter a faster decrease in work output was found in the youngest group (approximately 2% each contraction) compared to the older groups (approximately 0.7% each contraction). After 40 contractions the reduction in work output was significantly different only between the youngest group and the two oldest groups (-30% vs -5%). These differences in fatigue were not associated with differences in adenosine 5′-triphosphate and phosphocreatine concentrations or in lactate production. Total work output and high-energy phosphate consumption increased by approximately 555% and 380% from age 40 to 120 days, respectively. Consequently, efficiency was significantly higher (approximately 32%) in the older groups compared to 40-day-old animals. Normalized for muscle mass, mean rate of high-energy phosphate consumption was similar in all groups whereas mean power output was significantly lower in the youngest group (approximately 46%). Thus, the difference in efficiency between the young and the other groups may be attributed to a lower external power production in the youngest group rather than changes in energy turnover