The influence of muscle action on the acute growth hormone response to resistance exercise and short-term detraining.

The effects of resistance training with concentric or concentric-eccentric muscle actions on the acute hormonal response to a resistance exercise protocol was investigated. Thirty-two men completed a 19 week lower-body resistance training program (consisting of the leg press and leg extension exercises) in which they (1) performed concentric actions only (CON); (2) performed both concentric and eccentric actions (CON-ECC); (3) performed double concentric actions for each repetition (CON-CON); or (4) did not exercise. Following training each subject performed two exercise tests consisting of three sets of 30 isokinetic concentric (day 1) and eccentric (day 2) knee extensions separated by 48 h. The exercise tests were repeated following 4 weeks of detraining. Blood samples were obtained before and after each exercise test. Serum growth hormone (GH) was significantly (P< 0.05) greater for the concentric test in groups CON and CON-CON whereas GH was lower for the concentric test in CON-ECC compared with the eccentric test prior to detraining. Following detraining, GH was greater during the concentric test in CON-ECC than in the eccentric test, whereas no differences were observed between the concentric and eccentric tests in CON and CON-CON and the acute GH response to resistance exercise was attenuated. These data indicate that GH is sensitive to muscle action type with differential responses observed with resistance exercise after short-term detraining.

Vulnerability to dysfunction and muscle injury after unloading.

OBJECTIVE: To test whether unloading increases vulnerability to eccentric exercise-induced dysfunction and muscle injury. DESIGN: Before-after trial. SETTING: General community. PATIENTS OR OTHER PARTICIPANTS: Two women and 5 men (73 +/- 3kg [mean +/- SE]) who were active college students but were not trained in lower body resistance exercise volunteered. INTERVENTION: Five weeks of unilateral lower limb suspension (ULLS), which has been shown to decrease strength and size of the unloaded, left, but not load-bearing, right quadriceps femoris muscle group (QF) by 20% and 14%, respectively; performance of 10 sets of ten eccentric actions with each QF immediately after the ULLS strength tests with a load equivalent to 65% of the post-ULLS eccentric 1-repetition maximum. MAIN OUTCOME MEASURE(S): Concentric and eccentric 1-repetition maximum for the left, unloaded and the right, load-bearing QF measured immediately after ULLS and 1,4,7,9, and 11 days later; cross-sectional area and spin-spin relaxation time (T2) of each QF as determined by magnetic resonance imaging and measured the last day of ULLS and 3 days later. RESULTS: The mean load used for eccentric exercise was 23 +/- 2 and 30 +/- 3kg for the left, unloaded and right, load-bearing QF, respectively. The concentric and eccentric 1-repetition maximum for the unloaded and already weakened left QF was further decreased by 18% (p = .000) and 27% (p = .000), respectively, 1 day after eccentric exercise. Strength did not return to post-ULLS levels until 7 days of recovery. The right, load-bearing QF showed a 4% decrease (p = .002) in the eccentric 1-repetition maximum 1 day after eccentric exercise. The left, unloaded QF showed an increase in T2 (p = .002) in 18% of its cross-sectional area 3 days after the eccentric exercise, thus indicating muscle injury. The right, load-bearing QF showed no elevation in T2 (p = .280). CONCLUSION: Unloading increases vulnerability to eccentric exercise-induced dysfunction and muscle injury, even at relatively light loads.

Effect of short-term unweighting on human skeletal muscle strength and size.

This study examined the effect of short-term unweighting on strength and size of lower limb muscle groups to predict probable responses to planned 16-d Shuttle flights. Subjects were 10 healthy males, exposed to 16 d of lower limb suspension (LLS). All ambulatory activity was performed on crutches while wearing a shoe with a 10-cm thick sole on the right foot. This eliminated ground contact by the left foot, and thereby, body weight bearing by the left lower limb. Biopsies of the left vastus lateralis muscle and T1 weighted magnetic resonance (MR) images (1.5 Tesla, TR/TE 600/20, 4 nex, 48 cm rectangular FOV, 10 mm transaxial slices at 5-mm intervals) of both thighs were used to examine muscle morphology. The in vivo speed-torque relation for the left and right quadriceps femoris (QF) muscle group was used to measure strength. Fiber type composition and average cross-sectional area were not altered by LLS. The speed-torque relation for the left QF was down-shifted 12% (p < 0.05) after LLS. There was no effect of speed or type of muscle action (eccentric, isometric, or concentric). The speed-torque relation for the right QF showed no change after LLS. Average cross-sectional area of the left QF in eight MR images of the mid-thigh decreased (p < 0.05) 8% with LLS (70 +/- 3 to 64 +/- 4 cm2), while the right QF showed no change (72 +/- 4 to 72 +/- 4 cm2). The hamstring muscle group showed no change in average cross-sectional area after LLS.(ABSTRACT TRUNCATED AT 250 WORDS)

Skeletal muscle myosin heavy chain composition and resistance training.

We recently reported that 19 wk of heavy resistance training caused a decrease in the percentage of type IIb and an increase in the percentage of type IIa fibers as determined by qualitative histochemical analyses of myofibrillar adenosinetriphosphatase activity of biopsies of musculus vastus lateralis (Hather et al. Acta Physiol. Scand. 143: 177-185, 1991). These data were interpreted to suggest that resistance training had caused transformation among the fast-twitch fiber subtypes. To more clearly establish the influence of resistance training on muscle fiber composition, biopsies from the original study were analyzed biochemically for myosin heavy chain (MHC) composition by use of sodium dodecyl sulfate-polyacrylamide gel electrophoresis and histochemically for fiber types by use of myofibrillar adenosinetriphosphatase activity. The results show that after training (n = 13), IIb MHC composition decreased (P < 0.05) from 19 +/- 4 to 7 +/- 1%. IIa MHC, in contrast, increased (P < 0.05) from 48 +/- 3 to 60 +/- 2%. These responses were essentially mirrored by alterations in fiber type distribution. The percentage of type IIb fibers decreased (P < 0.05) from 18 +/- 3 to 1 +/- 1%, whereas the percentage of type IIa fibers increased from 46 +/- 4 to 60 +/- 3% (P < 0.05). Neither I MHC composition nor type I fiber percentage changed with training. The control group (n = 4) showed no changes in MHC composition or fiber type distribution. These results suggest that heavy resistance training alters MHC composition in human skeletal muscle, presumably reflecting a change in genetic expression.

Skeletal muscle responses to unloading with special reference to man.

The limited space flight data suggest that exposure to microgravity decreases muscle strength in humans and muscle mass in lower mammals. Several earth-based models have been used to address the effect of unloading on the human neuromuscular system due to the limited access of biological research to long-term space flight. Bedrest eliminates body weight bearing of both lower limbs. Unilateral lower limb suspension (ULLS), where all ambulatory activity is performed on crutches with an elevated sole on the shoe of one foot, has recently been used to unload one lower limb. The results from studies using these two models support their efficacy. The decrease in strength of m. quadriceps femoris, for example, after four to six weeks of bedrest, ULLS or space flight is 20 to 25%. The results from the earth-based studies show that this response can be attributed in part to a decrease in the cross-sectional area of the KE which reflects muscle fiber atrophy. The results from the ground based studies also support the limited flight data and show that reductions in strength are larger in lower than upper limbs and in extensor than flexor muscle groups. They also raise issue with the generally held concept that postural muscle is most affected by unweighting. Slow-twitch fibers in lower limb muscles of mixed fiber type composition and muscle composed mainly of slow-twitch fibers do not preferentially atrophy after bedrest or ULLS. Taken together, the data suggest that unloading causes remarkable adaptations in the neuromuscular system of humans. It should be appreciated, however, that this area of research is in its infancy.

Skeletal muscle responses to lower limb suspension in humans.

Eight subjects participated in a 6-wk unilateral lower limb suspension (ULLS) study to determine the influence of reduced weight bearing on human skeletal muscle morphology. The right shoe was outfitted with a platform sole that prevented the left foot from bearing weight while walking with crutches, yet it allowed freedom of movement about the ankle, knee, and hip. Magnetic resonance images pre- and post-ULLS showed that thigh muscle cross-sectional area (CSA) decreased (P less than 0.05) 12% in the suspended left lower limb, whereas right thigh muscle CSA did not change. Likewise, magnetic resonance images collected post-ULLS showed that muscle CSA was 14% smaller (P less than 0.05) in the left than in the right leg. The decrease in muscle CSA of the thigh was due to a twofold greater response of the knee extensors (-16%, P less than 0.05) than knee flexors (-7%, P less than 0.05). The rectus femoris muscle of the knee extensors showed no change in CSA, whereas the three vastus muscles showed similar decreases of approximately 16% (P less than 0.05). The apparent atrophy in the leg was due mainly to reductions in CSA of the soleus (-17%) and gastrocnemius muscles (-26%). Biopsies of the left vastus lateralis pre- and post-ULLS showed a 14% decrease (P less than 0.05) in average fiber CSA. The decrease was evident in both type I (-12%) and II (-15%) fibers. The number of capillaries surrounding the different fiber types was unchanged after ULLS.(ABSTRACT TRUNCATED AT 250 WORDS)

Influence of eccentric actions on skeletal muscle adaptations to resistance training.

Three different training regimens were performed to study the influence of eccentric muscle actions on skeletal muscle adaptive responses to heavy resistance exercise. Middle-aged males performed the leg press and leg extension exercises two days each week. The resistance was selected to induce failure within six to twelve repetitions of each set. Group CON/ECC (n = 8) performed coupled concentric and eccentric actions while group CON (n = 8) used concentric actions only. They did four or five sets of each exercise. Group CON/CON (n = 10) performed twice as many sets with only concentric actions. Eight subjects did not train and served as controls. Tissue samples were obtained from m. vastus lateralis using the biopsy technique before and after 19 weeks of training, and after four weeks of detraining. Histochemical analyses were performed to assess fibre type composition, fibre area and capillarization. Training increased (P less than 0.05) Type IIA and decreased (P less than 0.05) Type IIB fibre percentage. Only group CON/ECC increased Type I area (14%, P less than 0.05). Type II area increased (P less than 0.05) 32 and 27%, respectively, in groups CON/ECC and CON/CON, but not in group CON. Mean fibre area increased (P less than 0.05) 25 and 20% in groups CON/ECC and CON/CON, respectively. Capillaries per fibre increased (P less than 0.05) equally for Type I and Type II fibres. Capillaries per fibre area for both fibre types, however, increased (P less than 0.05) only in groups CON and CON/CON. The changes in fibre type composition and capillary frequency were manifest after detraining.(ABSTRACT TRUNCATED AT 250 WORDS)

Histochemical demonstration of skeletal muscle fibre types and capillaries on the same transverse section.

Serial transverse sections of m. vastus lateralis biopsies from six healthy men were reacted: (1) for myofibrillar adenosine triphosphatase (mATPase) to identify fibre types; or, (2) with alpha-amylase, periodic acid-Schiff (alpha PAS) to visualize capillaries. Sections were also processed with a new histochemical method for identification of fibre types and capillaries on the same skeletal muscle slice (mATPase/alpha PAS). Fibre type composition using either method was 41% type I, 37% type IIA and 22% type IIB. Types I, IIA and IIB least diameter areas (mean +/- SE, micron2) were similar in sections processed for mATPase/alpha PAS or mATPase (3976 +/- 338, 5187 +/- 373 and 4389 +/- 514 vs. 4092 +/- 345, 5100 +/- 360 and 4289 +/- 474, respectively). The number of capillaries per mm2 and per fibre did not differ in sections processed using the alpha PAS (315 +/- 29 and 1.48 +/- 0.12) or mATPase/alpha PAS (317 +/- 25 and 1.43 +/- 0.10) method. The number of capillaries was greater (P less than 0.05) around types I or IIA than type IIB fibres whether a section was processed for mATPase/alpha PAS (4.5 +/- 0.2 or 4.6 +/- 0.2 vs. 3.4 +/- 0.3) or when fibre profiles of serial sections reacted for mATPase or alpha PAS were 'matched' (4.5 +/- 0.2 or 4.4 +/- 0.2 vs. 3.4 +/- 0.3). The results indicate that histochemical and morphometric measures can be made on the same transverse section using the new method with substantial savings of time, cost and energy.

Effect of voluntary vs. artificial activation on the relationship of muscle torque to speed.

The speed-torque relationship of the right knee extensor muscle group was investigated in eight untrained subjects (28 +/- 2 yr old). Torque was measured at a specific knee angle during isokinetic concentric or eccentric actions at nine angular velocities (0.17-3.66 rad/s) and during isometric actions. Activation was by "maximal" voluntary effort or by transcutaneous tetanic electrical stimulation that induced an isometric torque equal to 60% (STIM 1) or 45% (STIM 2) of the voluntary isometric value. Torque increased (P less than 0.05) to 1.4 times isometric as the speed of eccentric actions increased to 1.57 rad/s for STIM 1 and STIM 2. Thereafter, increases in eccentric speed did not further increase torque. Torque did not increase (P greater than 0.05) above isometric for voluntary eccentric actions. As the speed of concentric actions increased from 0.00 to 3.66 rad/s, torque decreased (P less than 0.05) more (P less than 0.05) for both STIM 1 and STIM 2 (two-thirds) than for voluntary activation (one-half). As a result of these responses, torque changed three times as much (P less than 0.05) across speeds of concentric and eccentric actions with artificial (3.4-fold) than voluntary (1.1-fold) activation. The results indicate that with artificial activation the normalized speed-torque relationship of the knee extensors in situ is remarkably similar to that of isolated muscle. The relationship for voluntary activation, in contrast, suggests that the ability of the central nervous system to activate the knee extensors during maximal efforts depends on the speed and type of muscle action performed.

Force and EMG signal patterns during repeated bouts of concentric or eccentric muscle actions.

Healthy males (n = 14) performed three bouts of 32 unilateral, maximal voluntary concentric (CON) or eccentric (ECC) quadriceps muscle actions on separate days. Surface electromyography (EMG) of the m. vastus lateralis (VL) and m. rectus femoris (RF) and torque were measured. Integrated EMG (IEMG), mean (MPF) and median power frequencies and torque were averaged for seven separate blocks of four consecutive muscle actions. Torque was greater (P less than 0.05) for ECC than for CON muscle actions at the start of exercise. It did not decline throughout ECC exercise, but decreased (P less than 0.05) markedly for each bout and over bouts of CON exercise. Thus, torque overall was substantially greater (P less than 0.05) for ECC than for CON exercise. At the start of exercise IEMG of VL or RF was greater (P less than 0.05) for CON than for ECC muscle actions. This was also true for overall IEMG activity during exercise. The IEMG increased (P less than 0.05) modestly for both muscles during each bout of CON or ECC muscle actions, but did not change for the VL over bouts. The IEMG of RF decreased (P less than 0.05) modestly over CON but not ECC exercise bouts. At the beginning of the first bout of exercise the IEMG/torque ratio was twofold greater (P less than 0.05) for CON than ECC muscle actions. The ratio of IEMG/torque increased (P less than 0.05) markedly during CON but did not change during ECC exercise. Thus, by the end of the third bout there was a fivefold difference (P less than 0.05).(ABSTRACT TRUNCATED AT 250 WORDS)

Properties of standard avian slow muscle grafts following long-term regeneration.

The tonic anterior latissimus dorsi (ALD) of adult pigeons was orthotopically homografted and evaluated after 11 months of regeneration for histological, histochemical, electromyographic (EMG), and mechanical properties. The resting EMG activity of the grafts was lower in amplitude than that of the controls, but showed the tonic pattern typical for these tonic muscles. The control and grafted muscles had a histochemically homogeneous population of fibers with moderate myofibrillar adenosine triphosphatase activity. Succinic dehydrogenase activity was moderate for the control muscles, but low for the grafts. The regenerated muscles had fewer and smaller fibers and had much larger intersynaptic distances. Both the regenerated and the contralateral control muscles were slow contracting and maintained tetanic tension for prolonged periods with direct electrical stimulation. The relaxation was slower in the grafted muscle than in the control. The grafts produced 40% of the maximum tension of the control muscles, but the rate of tension development was similar between the two groups. The results indicate that the tonic properties were regenerated, but the innervation pattern was altered and the grafted muscles did not have normal mature fibers even after long-term regeneration.

Quantification and origin of spindles in orthotopic and heterotopic grafts of avian muscles.

Regeneration of muscle spindles was quantified in a series of orthotopically and heterotopically autografted muscles of pigeons. Significantly fewer spindles relative to numbers of extrafusal fibers were present in grafts than in normal muscles. These results are in marked contrast to observations of free-grafted muscles of rats. A majority of grafts of the metapatagialis, a muscle devoid of spindles, into the site of the anterior latissimus dorsi contained spindles. A few spindles were present in grafts of the extensor digitorum communis, which normally contains many spindles, into the site formerly occupied by the metapatagialis whereas muscle spindles were absent in orthotopic grafts of the metapatagialis muscle. These observations suggest that the spindle-like structures observed in the extensor digitorum communis muscles, which regenerated in the sites of the metapatagialis, were derived from spindles of the donor muscle. Thus muscle spindles in transplanted avian muscle can form by two distinct developmental processes.