Capillary supply of the quadriceps femoris muscle of man: adaptive response to exercise

1. Five subjects trained for 8 weeks on a bicycle ergometer for an average of 40 min/day, four times a week at a work load requiring 80% of the maximal oxygen uptake (V(O2 max.)). V(O2 max.) determinations were performed, and muscle biopsies from the quadriceps femoris muscle (vastus lateralis) were taken before, as well as repeatedly during, the training period. The muscle biopsies were histochemically stained for fibre-types (myofibrillar ATPase) and capillaries (amylase-PAS method), and analysed biochemically for succinate dehydrogenase and cytochrome oxidase activities.2. The training programme resulted in a 16% increase in V(O2 max.), a 20% increase in capillary density, a 20% increase in mean fibre area, and an approximately 40% increase in the activities of succinate dehydrogenase and cytochrome oxidase.3. The capillary supply to type I, IIA and IIB fibres, expressed as the mean number of capillaries in contact with each fibre-type, relative to fibre-type area, increased equally.4. The present study shows that endurance training constitutes a powerful stimulus for capillary proliferation in human skeletal muscle.     

Adaptive changes in work capacity,skeletal muscle capillarization and enzyme levels during training and detraining

Six male subjects exercised on a bicycle ergometer 30 min with left leg and 30 min with right leg 3 times a week for 8 weeks. This training resulted in a 14.6% increase in V?_o2 max with two-leg exercise and a 23.1% increase with one-leg exercise. A significant decrease towards pretraining V?_o2 max was seen during the following 8 weeks of detraining. Muscle biopsy samples were obtained at rest from m. vastus lateralis before and after training and 4 and 8 weeks after training. During training the number of capillaries per mm² and the number of capillaries per fiber increased about 20%. The number of capillaries around each fibre type (CA) increased 20–30%. The average area of each fibre type increased only about 5%. The fibre area per CA decreased by about 10%. During 8 weeks of detraining decreases were seen in the number of capillaries per fibre, CA and in fibre area, while fibre area per CA and number of capillaries per mm² were almost unchanged at the end of the detraining period. Pronounced increases in activities of oxidative enzymes were observed after training, while only minor increases were seen in glycolytic enzyme activities. All enzyme activities decreased towards pre-training levels during detraining. The results indicate that the training-induced improvement in oxidative capacity and in muscle capillarization expressed as capillaries per fibre and CA disappears within 8 weeks after cessation of training. However, the fibre area per CA and number of capillaries per mm² point at a favourable long term effect on the average diffusion distance between capillaries and muscle fibres.     

Muscle Adaptation to Extreme Endurance Training in Man

To evaluate the effect of extreme endurance training on muscle fibre composition and activities of oxidative enzymes in different fibre types biopsies were taken from vastus lateralis, gastrocnemius and deltoideus of elite orienteers. Comparisons were made between the (trained) leg muscles and the (relatively untrained) arm muscles, and with leg muscles of 16–18 years old boys. The orienteers had the same percentage type I fibres in vastus lateralis and gastrocnemius as in deltoideus, but higher percentage type I fibres in vastus lateralis compared with the controls. The similarity between trained and untrained muscle in the orienteers suggests that training had not caused the high percentage type I fibres which rather might be the result of selection of individuals with the best prerequisites for high oxidative capacity. However, the distribution of type II subgroups in the leg muscles of the orienteers differed from both their own deltoideus and leg muscles of the controls, the relationship IIA/IIB being altered in favour of the more oxidative IIA. The leg muscles of the orienteers also showed an increased occurrence of the normally rare IIC fibre. These latter findings point at the possibility of a training induced alteration in the subgroup pattern. Unlike in the controls there was no significant difference in succinate dehydrogenase activity, measured in single fibres, between type I and II fibres in gastrocnemius of the orienteers. Thus, type II fibres have the ability metabolically to adapt to high oxidative demands. This might to some extent be mediated by a conversion from IIB to IIA form.     

Physical training in obese women

Summary Peripheral adaptations to 3 months of physical endurance training without food restrictions were studied in skeletal muscles of 14, middle-aged, physically untrained, obese women.In comparison to aged-matched controls of normal weight, the obese group showed significantly lower isometric endurance. In the obese group, physical training resulted in a significant increase of maximal isometric and isokinetic strength. Isokinetic but not isometric endurance also increased after training. The isometric strength of obese women showed a positive correlation with the percentage of FTb fibres.The training (50 min/day, 3 days/w) did not result in any change in body weight, body fat, and the number and weight of fat cells. The 20% increase of after training was found to be significantly correlated with the increase in the number of capillaries around muscle fibres. The relative percentage of FTa fibres, the number of capillaries per fibre as well as the activities of citrate synthase, 3-hydroxy-acyl-CoA-dehydrogenase, and hexokinase showed a significant increase after training.The concentrations of glucose during OGTT showed a trend to decrease with a significant decrease at the end glucose curve (120-min value). The concentration of insulin and C peptide and the insulin removal did not change after training. The changes in the concentration of glucose during OGTT was significantly correlated with the increase in muscle capillarization and of dynamic endurance.     

The GLUT4 density in slow fibres is not increased in athletes. How does training increase the GLUT4 pool originating from slow fibres?

The influence of training on GLUT4 expression in slow- and fast-twitch skeletal muscle fibres was studied in male endurance-trained athletes and control subjects. The trained state was ensured by elevated maximal oxygen uptake (29%), as well as citrate synthase (60%) and 3-hydroxy-acyl-CoA dehydrogenase (38%) activities in muscle biopsy samples of the vastus lateralis. GLUT4 densities in slow- and fast-twitch fibres were measured by the use of a newly developed, sensitive method combining immunohistochemistry with morphometry, and no effect of training was found. GLUT4 density was higher in slow-twitch fibres compared to fast-twitch fibres (P<0.05) when biopsy samples from untrained subjects were examined. In athletes GLUT4 density was identical in slow- and fast-twitch fibres. Slow-twitch fibre diameters were 10% larger in the athletes (P<0.01), and slow-twitch fibre fractions were 140% of the fraction in the control group. Thus, GLUT4 originating from slow-twitch fibres was increased by 30% (P<0.02) in athletes. We conclude that long-lasting endurance training increases slow-twitch fibre GLUT4 expression by means of an elevated slow-twitch fibre mass in human skeletal muscle.     

Different responses of skeletal muscle following sprint training in men and women

Six male and ten female physically active students performed 30-s sprint training on a cycle ergometer three times a week for 4 weeks, to investigate the training responses of skeletal muscle and to evaluate possible sex differences in this respect. Three repeated sprint tests with a 20-min rest in between were performed and muscle biopsies from the vastus lateralis muscle were taken before and after the training period. Mean power (average of sprint I–III) and type HB cross-sectional fibre area increased only in the women (7% and 25% respectively) following sprint training, resulting in a decreased sex difference. There was an increase in total lactate dehydrogenase (LD) activity following sprint training in both sexes, although the levels were lower in the women both before and after training. Glycogen content increased and the activity of LD iso-enzyme I decreased in the women, but not in men. It was hypothesised that both the smaller areas of type II fibres and lower activity of LD generally seen in women may be due in part to less frequent activation of type TI fibres in women than in men. If this were the case, the women should respond to sprint training (a type of training that activates type II fibres) to a relatively greater extent than men. That the observed increase in type IIB fibre area in response to sprint training was greater in the women than in men supported the hypothesis of the study. However, the results for LD activity, which showed a similar response in the men and the women, did not support the hypothesis.     

Muscle adaptation to extreme endurance training in man.

To evaluate the effect of extreme endurance training on muscle fibre composition and activities of oxidative enzymes in different fibre types biopsies were taken from vastus lateralis, gastrocnemius and deltoideus of elite orienteers. Comparisons were made between the (trained) leg muscles and the (relatively untrained) arm muscles, and with leg muscles of 16--18 years old boys. The orienteers had the same percentage type I fibres and vastus lateralis and gastrocnemius as in deltoideus, but higher percentage type I fibres in vastus lateralis compared with the controls. The similarity between trained and untrained muscle in the orienteers suggests that training had not caused the high percentage type I fibres which rather might be the result of selection of individuals with the best prerequisites for high oxidative capacity. However, the distribution of type II subgroups in the leg muscles of the orienteers differed from both their own deltoideus and leg muscles of the controls, the relationship IIA/IIB being altered in favour of the more oxidative IIA. The leg muscles of the orienteers also showed an increased occurrence of the normally IIC fibre. These latter findings point at the possibility of a training induced alteration in the subgroup pattern. Unlike in the controls there was no significant difference in succinate dehydrogenase activity, measured in single fibres, between type I and II fibres in gastrocnemius of the orienteers. Thus, type II fibres have the ability metabolically to adapt to high oxidative demands. This might to some extent be mediated by a conversion from IIB to IIA form.     

Histochemical and biochemical changes in human skeletal muscle with age in sedentary males, age 22--65 years.

Biopsies for histochemical and biochemical analyses were taken from the vastus lateralis muscle of 55 untrained, healthy male subjects from 22 to 65 years of age. Fibre type distribution changed towards a decrease in the percentage of type II fibres, both in type IIA and type IIB fibres, whereas type IIB/IIA fibre ratio and type IIC percentage did not change with increasing age. It was found that the type IIB/IIA fibre ratio was inversely related to type I fibres, i.e. subjects rich in type I fibres had a relatively smaller proportion of type IIB fibres. Fibre area determinations revealed a selective decrease in type II fibre area. Consequently, the type II/I fibre area ratio and relative type II fibre area decreased. No changes in the specific activities of Mg2+ stimulated ATPase and myokinase were observed, while the activity of lactate dehydrogenase (LDH) was higher in the youngest groups than in the oldest. LDH isozyme pattern shifted towards a decrease in percentage distribution of the muscle specific isozymes and a corresponding decrease in muscle specific activity while the activity of the heart specific isozymes did not change.     

Effect of exercise training on muscle glucose transporter 4 protein and intramuscular lipid content in elderly men with impaired glucose tolerance

This study determined the effects of exercise training on adaptations of skeletal muscle including fibre composition, capillarity, intra-muscular triglyceride concentration (IMTG), as well as glucose transporter 4 protein (GLUT4) and metabolic enzyme activities. Percutaneous muscle biopsies from the vastus lateralis muscle were obtained from non-obese elderly Korean men (n=10; age range 58–67 years) with impaired glucose tolerance. Subjects performed 12 weeks of endurance exercise training (60–70% of the heart rate reserve). The training program improved the total GLUT4 protein expression (P<0.01), decreased the IMTG, increased the fatty acid oxidation capacity, and the number of capillaries around type 1 fibres (P<0.05), whereas no significant alteration was observed around type II fibres. All data are presented as the means together with the standard deviation. The results suggest that endurance training evokes morphological and biochemical changes in the skeletal muscle of elderly men with impaired glucose tolerance that may be considered to limit the development of type 2 diabetes.     

Myosin heavy chain isoform distribution in single fibres of bodybuilders

The purpose of this study was to investigate the long-term effects of high intensity resistance training on myosin heavy chain (MHC) isoform composition of single fibres. Muscle biopsies were obtained from the right vastus lateralis of eight bodybuilders (BB) and seven physical education students (PES). Histochemical analyses were used to determine the fibre type distribution and the fibre cross-sectional area. MHC isoform composition of single fibres was determined with protein electrophoresis. The percentage of fibres expressing MHC IIA and MHC I/IIA was larger in BB (P < 0.05), while MHC IIX was completely absent (P < 0.05). In contrast, myofibrilar ATPase histochemistry only revealed a significantly lower percentage of type IIX fibres in BB (P < 0.05). The muscle fibre profile in the vastus lateralis muscle of BB may represent an adaptation based on the mechanical and biochemical demands of the long-term resistance training.     

Histochemical and biochemical changes in human skeletal muscle with age in sedentary males,age 22–65 years

Biopsies for histochemical and biochemical analyses were taken from the vastus lateralis muscle of 55 untrained, healthy male subjects from 22 to 65 years of age. Fibre type distribution changed towards a decrease in the percentage of type II fibres, both in type IIA and type IIB fibres, whereas type IIB/IIA fibre ratio and type IIC percentage did not change with increasing age. It was found that the type IIB/flA fibre ratio was inversely related to type I fibres, i.e. subjects rich in type I fibres had a relatively smaller proportion of type IIB fibres. Fibre area determinations revealed a selective decrease in type II fibre area. Consequently, the type II/I fibre area ratio and relative type II fibre area decreased. No changes in the specific activities of Mg²⁺ stimulated ATPase and myokinase were observed, while the activity of lactate dehydrogenase (LDH) was higher in the youngest groups than in the oldest. LDH isozyme pattern shifted towards a decrease in percentage distribution of the muscle specific isozymes and a corresponding decrease in muscle specific activity while the activity of the heart specific isozymes did not change.     

The effect of strength training on estimates of mitochondrial density and distribution throughout muscle fibres

The purpose of this study was to investigate the effect of strength training (12 weeks, 3 days/week, four lower-body exercises) of young individuals (mean age 23.6 years) on estimates of mitochondrial distribution throughout muscle fibres. A control group (mean age 21.7 years) was followed simultaneously. Skeletal muscle biopsy samples were obtained from the vastus lateralis, pre- and post-training. The regional distribution of subsarcolemmal and intermyofibrillar mitochondrial populations was determined using quantitative histochemical staining of succinate dehydrogenase (SDH) in type I and II muscle fibres. Strength training resulted in significant increases of 26% and 28% in the cross-sectional area of type I and II fibres, respectively (P?P?相似文献   

Long-term resistance training improves force and unloaded shortening velocity of single muscle fibres of elderly women

Four elderly women (78 ± 4.3 years) were resistance trained (RT) for one year and needle biopsies of the vastus lateralis muscle were taken pre- and post-RT. A training intensity of 60% one-repetition maximum (1RM) was adopted for the entire duration of the study. Following RT, significant gain in isometric force of the quadriceps muscles was observed and MHC isoform distribution of vastus lateralis muscle showed a very significant shift from MHC-1 and MHC-2X towards MHC-2A, i.e. a bi-directional shift. A large population (n = 310) of individual skinned muscle fibres were dissected from biopsy samples and used for determination of cross-sectional area (CSA), specific force (Po/CSA) and unloaded shortening velocity (Vo) at 12°C and maximum activation. Fibres were classified on the basis of their MHC isoform content determined by SDS-PAGE in three pure fibre types (1, 2A and 2X) and two hybrid fibre types (1-2A and 2AX). Whereas the CSA of individual muscle fibres did not change, Po/CSA was significantly higher in both type 1 and type 2A fibres post training. Vo of type 1 fibres was significantly higher post-training, whereas no change in Vo was observed in type 2A fibres. The number of pure type 2X fibres was very low especially post-training and did not enable a statistically significant comparison. The data suggest that in elderly women moderate and prolonged RT: (1) can determine similar adaptations in MHC distribution of skeletal muscle to those expected in young subjects; (2) can improve function of muscle fibres.     

Muscle fibre types and enzyme activities after training with local leg ischaemia in man

Eight healthy men performed supine one-legged training on a bicycle ergometer 45 min per leg four times per week for 4 week. The ergometer and lower body were inside a pressure chamber, the opening of which was sealed at the level of the crotch. One leg trained with impeded leg blood flow (I-leg), induced by an increased (50 mmHg) chamber pressure, at the highest tolerable intensity. The contralateral leg trained at the same power under normal pressure (N-leg). Before and after training biopsies were taken from the vastus lateralis of both legs and maximal one-legged exercise tests were executed with both legs. Biopsies were repeated when the subjects had been back to their habitual physical activity for 3 months. Training increased exercise time to exhaustion, but more in the I-leg than in the N-leg. After training, the I-leg had higher activity of citrate synthase (CS), a marker of oxidative capacity, and lower activity of the M-subunit of lactate dehydrogenase isoenzymes. It also had a higher percentage of type-I fibres and a lower percentage of IIB fibres, larger areas of all fibre types and a greater number of capillaries per fibre. It is concluded that ischaemic training changes the muscle metabolic profile in a direction facilitating aerobic metabolism. An altered fibre-type composition may contribute, but is not enough prerequisite for the change.     

Distribution of different fibre types in human skeletal muscles 2. A study of cross-sections of whole m. vastus lateralis

In order to determine the total number of fibres and the extent to which the relative occurrence of different fibre types varies within m. vastus lateralis, 15 μm thick cross-sections of whole muscles were prepared. The total number of type 1 and type 2 fibres was determined in every 48th square millimetre of the section, and the results thus obtained were analysed using a computer program allowing an assessment of bivariate data in the form of contour plots. The total number of fibres varied both in proximal to distal direction in the same muscle and between individuals. No obvious correlation existed between the mean fibre area and the muscle cross-sectional area. The proportion of type 1 fibres in the whole muscle varied between individuals (from 44% to 57%) with a mean value for all five of 52%. The distribution of different fibre types varied within the muscle, mainly as a function of depth, with a predominance to type 2 fibres at the surface and type 1 fibres in deeper regions of the muscle. Thus, the fibre type distribution in m. vastus lateralis is not random. This must be taken into consideration when data on fibre type composition are compared with functional variables.     

High proportion of type I fibres in thigh muscle of young dancers

A previous study showed that adult female dancers have a high percentage of type I fibres in vastus lateralis. similar to that of endurance-trained female runners ar female cross-country skiers. It is not known if dancers already at an early age are characterized by a high percentage of type I fibres or develop a high percentage of type I fibres as a consequence of dance training. Furthermore. the muscle fibre composition of male dancers has not previously been studied. Therefore the aim of the study was to analyse skeletal muscle fibre characteristics in 10-year-old and 20-year-old dancers of both sexes. Age-matched boys and girls whose physical activity was average for their age groups served as controls. Muscle biopsies for histochemical analysis were obtained from vastus lateralis using the percutaneous needle technique. The major finding of the present study was that the vastus lateralis of young dancers of both sexes had a higher percentage of type I fibres than that of controls. Moreover. the higher type I percentage was seen not only in 20 year aids. but also in 10 year aids. who had begun their dance training at a professional level only a few weeks earlier. No significant difference in this respect was found between female and male dancers. In conclusion. the muscle fibre type composition in young dancers of both sexes differs from that of the average individual of the same age and is characterized by a high percentage of type I fibres.     

Muscle fibre type distribution,muscle cross-sectional area and maximal voluntary strength in humans

The relationship between maximum voluntary concentric strength, muscle fibre type distribution and muscle cross-sectional areas were examined in 23 subjects (7 female and 11 male phys. ed. students as well as 5 male bodybuilders). Maximal knee and elbow extension as well as elbow flexion torque at the angular velocities 30, 90 and 180 degrees per second was measured. Muscle biopsies were taken from vastus lateralis and m. triceps brachii. The muscle cross-sectional area of the thigh and upper arm was measured with computed tomography scanning. The maximal torque correlated strongly to the muscle cross-sectional area times an approximative measure on the lever arm (body height). Maximal tension developed per unit of muscle cross-sectional area did not correlate significantly with per cent type I fibre area and did not differ between the female and male students or bodybuilders. Neither did the relative decrease in torque with increasing contraction velocity show any significant relationship to the per cent type I fibre area. The total number of muscle fibres was estimated by dividing the muscle cross-sectional area with the mean fibre area of m. triceps brachii. The number of fibres did not seem to differ between the sexes.     

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)     

Effect of Strength Training on EMG of Human Skeletal Muscle

The effects of an 8 weeks period of systematic progressive strength training on the EMG activity of the leg extensor muscles (vastus lateralis and rectus fernoris) were investigated in 8 healthy male subjects (22–31 yrs). After training there were indications (n.s.) of a decline in integrated EMG (IEMG) during maximal isometric knee extension as well as in the IEMG vs isometric force relationship. The averaged motor unit potential (AMUP) did not demonstrate any significant changes due to the strength training regimen. In conformity with earlier findings no or only minor alterations were observed in anthropometrics, muscle enzyme activities and fibre composition. The fibre area ratio indicated a specific effect of the training stimuli on the fast twitch muscle fibres. Thus, EMG-analyses, as employed in the present study, did not provide any conclusive additional explanation as to the mechanisms behind the well established gains in niuscle strength performance induced by the applied strength training program.     

Muscle fibre size and capillarity in Korean diving women

AIM: Effects of prolonged habitual cold-water immersion on fibre size and capillarity in vastus lateralis muscle were studied in human beings. The hypothesis tested in the present study was that cold acclimatized human skeletal muscle would have reduced muscle fibre size and higher capillarity, favouring the idea of efficacy of recruitment under cold environment. METHODS: Ten women breath-hold divers (BHDs) and 10 active women (controls CONs) participated in this study. Muscle biopsy was obtained from vastus lateralis and determined fibre type composition and capillary density. RESULTS: A major finding was that all BHDs revealed a markedly smaller cross-sectional area (CSA) in all fibre types than the CONs, or even than any other morphological data reported in previous investigations. Furthermore, mean CSA of type II fibre (range 1205-2766 microm2) was much smaller than type I fibre (2343-4327 microm2). The number of capillaries per fibre in different fibre types in the BHDs was higher than in the CONs (P < 0.001), and diffusional area was smaller in type II fibres than in type I fibres (P < 0.001). The BHDs and the CONs have similarity in the percentage of type I fibres, but type II fibre was predominant in both groups. Interestingly the proportion of type IIx fibre in the BHDs was higher (31%) than in the CONs (22%). No significant difference was found in the thigh circumference between the groups. CONCLUSION: The present study demonstrates that prolonged habitual cold-water immersion may induce a decrease in fibre size and an increase in capillarity in human skeletal muscle.