Ageing influences myonuclear domain size differently in fast and slow skeletal muscle of rats.

AIM: In multinucleated skeletal muscle, a myonuclear domain is the region of cytoplasm governed by one nucleus, and myofibres are mosaics of overlapping myonuclear domains. Association of ageing and myonuclear domain is important in the understanding of sarcopenia and with prevention or combating age-related muscle declines. This study examined the effects of age, fibre type and muscle on nucleo-cytoplasmic (N/C) relationships as reflecting myonuclear domain size. METHODS: The N/C was compared in fibre types of soleus and plantaris muscles from young (n = 6) and ageing (n = 8) male Fisher 344 rats. RESULTS: There were no significant differences in fibre type composition or cross-sectional area of the soleus across ages. The old soleus had significantly more myonuclei, resulting in a significantly smaller myonuclear domain size. The plantaris muscle showed a higher percentage of slow fibres in old compared with young fibres. There were no differences in the number of myonuclei or in myonuclear domain size between young and older animals. CONCLUSION: We found muscle-specific differences in the effects of ageing on myonuclear domain, possibly as a result of reduced efficiency of the myonuclei in the slow muscles.

Effects of high-intensity resistance training on untrained older men. I. Strength, cardiovascular, and metabolic responses.

Most resistance training studies of older subjects have emphasized low-intensity, short-term training programs that have concentrated on strength measurements. The purpose of this study was, in addition to the determination of strength, to assess intramuscular and transport factors that may be associated with strength increments. Eighteen untrained men ages 60-75 years volunteered for the study; 9 were randomly placed in the resistance-training group (RT), and the other half served as untrained (UT) or control subjects. RT subjects performed a 16-week high-intensity (85-90% 1 repetition maximum (RT]) resistance training program (2 x/wk) consisting of 3 sets each to failure (6-8 repetitions based on 1 RM of 3 exercises): leg press (LP), half squat (HS), and leg extension (LE) with 1-2 minutes rest between sets. Pre- and post- training strength was measured for the 3 training exercises using a 1 RM protocol. Body fat was calculated using a 3-site skinfold method. Biopsies from the vastus lateralis m. were obtained for fiber type composition, cross-sectional area, and capillarization measurements. Exercise metabolism, electrocardiography, and arterial blood pressure were observed continuously during a progressive treadmill test, and resting echocardiographic data were recorded for all subjects. Pre- and post-training venous blood samples were analyzed for serum lipids. Resistance training caused significant changes in the following comparisons: % fat decreased in the RT group by almost 3%, strength improved for all exercises: LE = + 50.4%, LP = + 72.3%, HS = + 83.5%; type IIB fibers decreased and IIA fibers increased; cross-sectional areas of all fiber types (I, IIA, IIB) increased significantly, and capillary to fiber ratio increased but not significantly. No differences were noted for ECG and echocardiographic data. The RT group significantly improved treadmill performance and VO2max. Pre- and post-training serum lipids improved but not significantly. No significant changes occurred in any pre- to post-tests for the UT group. The results show that skeletal muscle in older, untrained men will respond with significant strength gains accompanied by considerable increases in fiber size and capillary density. Maximal working capacity, VO2max, and serum lipid profiles also benefited from high-intensity resistance training, but no changes were observed for HR max, or maximal responses of arterial blood pressure. Older men may not only tolerate very high intensity work loads but will exhibit intramuscular, cardiovascular, and metabolic changes similar to younger subjects.

Effects of high-intensity resistance training on untrained older men. II. Muscle fiber characteristics and nucleo-cytoplasmic relationships.

During growth and repair of skeletal muscle fibers, satellite cells become activated, undergo mitosis, and a daughter nucleus becomes incorporated into the muscle fiber to increase myonuclear numbers. An increase in myonuclei appears to be required for this postnatal growth. This study examined whether muscle fibers of elderly men can hypertrophy with strength training and, if so, whether they have the capacity to incorporate nuclei into the fibers. The sarcoplasmic area associated with each myonucleus was calculated in nine elderly men before and after 16 weeks of strength training, and compared to nine elderly control men. Muscle fiber type changes and myosin heavy chain composition were also compared. All major fiber types (I, IIA, IIB) became significantly larger after training, and a transition of type IIB fibers to IIA occurred with training. The area occupied by each fiber type correlated with myosin heavy chain percentage, and both of these changed similarly with strength training. The cytoplasm-to-myonucleus ratio increased, but not significantly (p = .07), with muscle fiber hypertrophy. Number of myonuclei per fiber and myonuclei per unit length of muscle fiber increased, but not significantly. Cross-sectional areas of the muscle fibers in untrained elderly men were much smaller than in untrained young men (when compared with our earlier studies). Training increased the sizes of the elderly muscle fibers to that of the untrained young men. This hypertrophy of muscle fibers by 30% with training resulted in no change in the cytoplasm-to-myonucleus ratio. This suggests that the myonuclear population continues to adapt to growth stimuli in the elderly muscles.

Fiber type composition of the vastus lateralis muscle of young men and women.

This study presents data collected over the past 10 years on the muscle fiber type composition of the vastus lateralis muscle of young men and women. Biopsies were taken from the vastus lateralis muscle of 55 women (21.2+/-2.2 yr) and 95 men (21.5+/-2.4 yr) who had volunteered to participate in various research projects. Six fiber types (I, IC, IIC, IIA, IIAB, and IIB) were classified using mATPase histochemistry, and cross-sectional area was measured for the major fiber types (I, IIA, and IIB). Myosin heavy chain (MHC) content was determined electrophoretically on all of the samples from the men and on 26 samples from the women. With the exception of fiber Type IC, no significant differences were found between men and women for muscle fiber type distribution. The vastus lateralis muscle of both the men and women contained approximately 41% I, 1% IC, 1% IIC, 31% IIA, 6% IIAB, and 20% IIB. However, the cross-sectional area of all three major fiber types was larger for the men compared to the women. In addition, the Type IIA fibers were the largest for the men, whereas the Type I fibers tended to be the largest for the women. Therefore, gender differences were found with regard to the area occupied by each specific fiber type: IIA>I>IIB for the men and I>IIA>IIB for the women. These data establish normative values for the mATPase-based fiber type distribution and sizes in untrained young men and women.

Fiber type composition of four hindlimb muscles of adult Fisher 344 rats.

The limb and trunk muscles of adult rats express four myosin heavy chain (MHC) isoforms, one slow (MHCI) and three fast (MHCIIa, MHCIId, and MHCIIb). The distribution of these isoforms correlates with fiber types delineated using myofibrillar actomyosin adenosine triphosphatase (mATPase) histochemistry. For example, type I fibers express MHCI and fiber types IIA, IID, and IIB express MHCIIa, MHCIId, and MHC-IIb, respectively. Fibers containing only one MHC isoform have been termed "pure" fibers. Recent evidence suggests that a population of "hybrid" fibers exist in rat skeletal muscle which contain two MHC isoforms. The purpose of the present investigation was to document the entire range of histochemically defined "pure" and "hybrid" fiber types in untreated muscles of the young adult Fisher 344 rat hindlimb. The selected hindlimb muscles (soleus, tibialis anterior, extensor digitorum longus, and gastrocnemius muscles) were removed from 12 male rats and analyzed for muscle fiber type distribution, cross-sectional area, and MHC content. Care was taken to delineate eight fiber types (I, IC, IIC, IIA, IIAD, IID, IIDB, and IIB) using refined histochemical techniques. Hybrid fibers were found to make up a considerable portion of the muscles examined (a range of 8.8-17.8% of the total). The deep red portion of the gastrocnemius muscle contained the largest number of hybrid fibers, most of which were the fast types IIAD (8.5+/-2.8%) and IIDB (5.2+/-2.3%). In conclusion, hybrid fibers make up a considerable portion of normal rat limb musculature and are an important population that should not be ignored.

Comparison of soleus muscles from rats exposed to microgravity for 10 versus 14 days.

The effects of two different duration space-flights on the extent of atrophy, fiber type composition, and myosin heavy chain (MHC) content of rat soleus muscles were compared. Adult male Fisher rats (n=12) were aboard flight STS-57 and exposed to 10 days of microgravity and adult ovariectomized female Spraque-Dawley rats (n=12) were aboard flight STS-62 for 14 days. Soleus muscles were bilaterally removed from the flight and control animals and frozen for subsequent analyses. Muscle wet weights, fiber types (I, IC, IIC, and IIA), cross-sectional area, and MHC content were determined. Although a significant difference was found between the soleus wet weights of the two ground-based control groups, they were similar with regard to MHC content (ca 90% MHCI and ca 10% MHCIIa) and fiber type composition. Unloading of the muscles caused slow-to-fast transformations which included a decrease in the percentage of type I fibers and MHCI, an increase in fibers classified as type IC, and the expression of two fast myosin heavy chains not found in the control rat soleus muscles (MHCIId and MHCIIb). Although the amount of atrophy (ca 26%) and the extent of slow-to-fast transformation (decrease in the percentage of MHCI from 90% to 82.5%) in the soleus muscles were similar between the two spaceflights, the percentages of the fast MHCs differed. After 14 days of spaceflight, the percentage of MHCIIa was significantly lower and the percentages of MHCIId and MHCIIb were significantly higher than the corresponding MHC content of the soleus muscles from the 10-day animals. Indeed, MHCIId became the predominant fast MHC after 14 days in space. These data suggest fast-to-faster transformations continued during the longer spaceflight.

The effects of short-term resistance training on endocrine function in men and women.

This investigation examined hormonal adaptations to acute resistance exercise and determined whether training adaptations are observed within an 8-week period in untrained men and women. The protocol consisted of a 1-week pre-conditioning orientation phase followed by 8 weeks of heavy resistance training. Three lower-limb exercises for the quadriceps femoris muscle group (squat, leg press, knee extension) were performed twice a week (Monday and Friday) with every other Wednesday used for maximal dynamic 1 RM strength testing. Blood samples were obtained pre-exercise (Pre-Ex), immediately post-exercise (IP), and 5 min post-exercise (5-P) during the first week of training (T-1), after 6 weeks (T-2) and 8 weeks (T-3) of training to determine blood concentrations of whole-blood lactate (LAC), serum total testosterone (TT), sex-hormone binding globulin (SHBG), cortisol (CORT) and growth hormone (GH). Serum TT concentrations were significantly (P < or = 0.05) higher for men at all time points measured. Men did not demonstrate an increase due to exercise until T-2. An increase in pre-exercise concentrations of TT were observed both for men and women at T-2 and T-3. No differences were observed for CORT between men and women; increases in CORT above pre-exercise values were observed for men at all training phases and at T-2 and T-3 for women. A reduction in CORT concentrations at rest was observed both in men and women at T-3. Women demonstrated higher pre-exercise GH values than men at all training phases; no changes with training were observed for GH concentrations. Exercise-induced increases in GH above pre-exercise values were observed at all phases of training. Women demonstrated higher serum concentrations of SHBG at all time points. No exercise-induced increases were observed in men over the training period but women increased SHBG with exercise at T-3. SHBG concentrations in women were also significantly higher at T-2 and T-3 when compared to T-1 values. Increases in LAC concentrations due to exercise were observed both for men and women for all training phases but no significant differences were observed with training. These data illustrate that untrained individuals may exhibit early-phase endocrine adaptations during a resistance training program. These hormonal adaptations may influence and help to mediate other adaptations in the nervous system and muscle fibers, which have been shown to be very responsive in the early phase of strength adaptations with resistance training.

Myonuclear loss in atrophied soleus muscle fibers.

BACKGROUND: A skeletal muscle fiber consists of many successive "territories," each controlled by the nucleus residing in that territory. Because nuclei appear to control a specific amount of territory (nuclear domain), nuclei must be added to accommodate an increase in fiber size. Because growth and hypertrophy require the addition of nuclei to fibers, it is of interest to determine whether atrophy causes a decrease in myonuclear number. This study compared the myonuclear population in the soleus muscles of rats that had undergone atrophy due to 10 days of spaceflight in the space shuttle, Endeavour, with muscles of ground-based control animals (10 rats each). METHODS: Myofibrillar ATPase activity was used to determine the major skeletal muscle fiber types in control rats and those having spent 10 days in space, and dystrophin antibodies were used to label the sarcolemma to identify underlying myonuclei. RESULTS: Type I and II fibers were atrophied after the flight, but type I fibers were atrophied twice as much as type II. Myonuclei were counted in identified and measured fibers, and the distribution normalized to number per millimeter of fiber circumference; this was significantly greater in type II than in type I fibers in both groups of rats. However, although the muscle fibers from flight animals were significantly atrophied, the normalized number of nuclei were identical between control and flight animals, indicating that nuclei decreased in numbers as the fibers atrophied. CONCLUSION: The nuclear domain is under strict control, and a decrease in the domain, as induced by atrophy, will cause nuclear degeneration and loss, which maintains a relatively constant size of the nuclear domain.

Differential titin isoform expression in human skeletal muscle.

Mammalian skeletal muscle expresses at least two isoforms of the cytoskeletal protein titin (connectin; MW approximately 3000 kDa). These isoforms are associated with different passive force curves, and thus may affect physical performance. To study the distribution of titin and its possible influence on performance in humans, muscle biopsies were obtained from 15 males (mean +/- SE; age = 25.4 +/- 2.9 years, height = 177.7 +/- 1.8 cm, weight = 76.5 +/- 2.2 kg). Two biopsies were obtained on separate occasions from both the right and left vastus lateralis, and one biopsy each from the lateral head of the right gastrocnemius and the right soleus, with all biopsies handled identically. Fibre type analyses were performed via mATPase histochemistry. Expression of titin and myosin heavy chain isoforms were determined by SDS-PAGE. Titin bands in the resulting gels were highly repeatable and were verified by migration patterns, as well as Western blot analysis. Two groups of subjects were identified: group 1 (n = 10) expressed only one titin isoform (titin-1) in all biopsies, and group 2 (n = 5) expressed two titin isoforms (titin-1 and titin-2) in all biopsies. No significant differences (P > 0.05) between groups were observed for percentage fibre types, percentage fibre type areas, fibre type cross-sectional areas, and percentage myosin heavy chain expression when comparing individual muscles, sampling times or bilateral comparisons. This is the first report of differential titin isoform expression in healthy, mature human skeletal muscle, but it is not clear why this occurs or what influence this may have on performance.

Effects of bovine growth hormone analogs on mouse skeletal muscle structure.

Skeletal muscles of transgenic mice expressing altered bovine growth hormones (bGH) have been compared with those of nontransgenic mice to determine whether muscle fiber type-specific responses or histopathologies are associated with the altered gene. The slow soleus and predominantly fast gastrocnemius muscles were prepared for myofibrillar ATPase activity (to determine muscle fiber type) and histological examination from mice that were either giant (M4 line), larger than normal (M11 line), dwarf (G119K line), or nontransgenic (NTC). No histopathology was observed in any of the muscles. Although body weights were significantly different between all four lines of mice, only the giant M4 mice had significantly larger muscle fibers than the other lines of mice, while neither the G119K nor M11 lines were significantly different from the NTC for either muscle. No fiber type-specific differences were noted. These results suggest that the different muscles are the product of differences in numbers of muscle fibers expressed in the G119K and M11 lines of mice; the increase in body mass matched the fiber size growth only in the giant M4 line. Therefore, the altered bGH genes may be acting on fetal liver and myoblast/myotube GH receptors to change the GH and IGF-I regulated pattern of muscle development, and eventually, to determine the adult muscle fiber numbers.

Effects of sprint cycle training on human skeletal muscle.

Eleven men sprint trained two to three times per week for 6 wk to investigate possible exercise-induced slow-to-fast fiber type conversions. Six individuals served as controls. Both groups were tested at the beginning and end of the study to determine anaerobic performance and maximal oxygen consumption. In addition, pre- and postbiopsies were extracted from the vastus lateralis muscle and were analyzed for fiber type composition, cross-sectional area, and myosin heavy chain (MHC) content. No significant changes were found in anaerobic or aerobic performance variables for either group. Although a trend was found for a decrease in the percentage of type IIb fibers, high-intensity sprint cycle training caused no significant changes in the fiber type distribution or cross-sectional area. However, the training protocol did result in a significant decrease in MHC IIb with a concomitant increase in MHC IIa for the training men. These data appear to support previous investigations that have suggested exercise-induced adaptations within the fast fiber population (IIb-->IIa) after various types of training (endurance and strength).

Skeletal muscle adaptations during early phase of heavy-resistance training in men and women.

An 8-wk progressive resistance training program for the lower extremity was performed twice a week to investigate the time course for skeletal muscle adaptations in men and women. Maximal dynamic strength was tested biweekly. Muscle biopsies were extracted at the beginning and every 2 wk of the study from resistance-trained and from nontrained (control) subjects. The muscle samples were analyzed for fiber type composition, cross-sectional area, and myosin heavy chain content. In addition, fasting blood samples were measured for resting serum levels of testosterone, cortisol, and growth hormone. With the exception of the leg press for women (after 2 wk of training) and leg extension for men (after 6 wk of training), absolute and relative maximal dynamic strength was significantly increased after 4 wk of training for all three exercises (squat, leg press, and leg extension) in both sexes. Resistance training also caused a significant decrease in the percentage of type IIb fibers after 2 wk in women and 4 wk in men, an increase in the resting levels of serum testosterone after 4 wk in men, and a decrease in cortisol after 6 wk in men. No significant changes occurred over time for any of the other measured parameters for either sex. These data suggest that skeletal muscle adaptations that may contribute to strength gains of the lower extremity are similar for men and women during the early phase of resistance training and, with the exception of changes in the fast fiber type composition, that they occur gradually.

Muscle fiber types of women after resistance training--quantitative ultrastructure and enzyme activity.

Muscle biopsies of the vastus lateralis muscle taken before and after 18 weeks of resistance training were compared by preparing frozen cross sections for electron microscopy and using adjacent sections for fiber typing by myosin ATPase activity. Quantitative ultrastructural changes were observed in histochemically-identified muscle fiber types of twelve young women who underwent the training. The percentage of type IIB fibers decreased and IIA fibers increased. The cross-sectional area of all major fiber types increased with training. The absolute volume of myofibrils, intermyofibrillar space, and mitochondria increased with training for most major fiber types (type I, IIA and IIAB), but the relative volume percentages were not significantly changed because of corresponding fiber hypertrophy. Mean mitochondrial size for types I and IIA and myofibril size for types IIC and IIB increased significantly with training. The capillary number per fiber and density did not change with training. Activity levels were measured for selected glycolytic and oxidative enzymes. Cytochrome oxidase and hexokinase increased significantly with training, while creatine kinase, citrate synthase, phosphofructokinase, glyceraldehyde phosphate dehydrogenase and hydroxyacyl CoA dehydrogenase enzymes were not significantly altered. The results suggest that this type of high-repetition resistance training causes the intracellular components of all fiber types to increase proportionally with an increase in fiber size. In addition, the enzyme analysis indicates the muscle as a whole may increase its oxidative phosphorylation capacity in conjunction with the decreased percentage of type IIB fibers.

Histochemical, biochemical, and ultrastructural analyses of single human muscle fibers, with special reference to the C-fiber population.

A muscle biopsy from the vastus lateralis muscle of a strength-trained woman was found to contain an unusual fiber type composition and was analyzed by histochemical, biochemical, and ultrastructural techniques. Special attention was given to the C-fibers, which comprised over 15% of the total fiber number in the biopsy. The mATPase activity of the C-fibers remained stable to varying degrees over the pH range normally used for routine mATPase histochemistry. Although a continuum existed, the C-fibers were histochemically subdivided into three main fiber types: IC, IIC, and IIAC. The IC fibers were histochemically more similar to the Type I, the IIAC were more similar to the Type IIA, and the IIC were darkly stained throughout the pH range. Biochemical analysis revealed that all C-fibers coexpressed myosin heavy chains (MHC) I and IIa in variable ratios. The histochemical staining intensity correlated with the myosin heavy chain composition such that the Type IC fibers contained a greater ratio of MHCI/MHCIIa, the IIAC contained a greater ratio of MHCIIa/MHCI, and the Type IIC contained equal amounts of these two heavy chains. Ultrastructural data of the C-fiber population revealed an oxidative capacity between fiber Types I and IIA and suggested a range of mitochondrial volume percent from highest to lowest such that I greater than IC greater than IIC greater than IIA-C greater than IIA. Under physiological conditions, it appears that the IC fibers represent Type I fibers that additionally express some fast characteristics, whereas the Type IIAC are Type IIA fibers that additionally express some slow characteristics. Fibers expressing a 50:50 mixture of MHCI and MHCIIa (IIC fibers) were rarely found. It is not known whether C-fibers represent a distinct population between the fast- and slow-twitch fibers that is specifically adapted to a particular usage or whether they are transforming fibers in the process of going from fast to slow or slow to fast.

Assessment of skeletal muscle damage in successive biopsies from strength-trained and untrained men and women.

The effects of repeated biopsy sampling on muscle morphology was qualitatively and quantitatively assessed in strength-trained and untrained men and women. College-age men (13) and women (8) resistance trained twice a week for 8 weeks. A progressive resistance-training program was performed consisting of squats, leg presses, and leg extensions. Nontraining men (7) and women (5) served as controls. Muscle biopsy specimens and fasting bloods were obtained at the beginning and every 2 weeks and histochemical, biochemical, and ultrastructural methods were employed to assess the type and amount of damage. Except for a few scattered atrophic fibers in 2 of the 33 biopsy samples, all initial specimens were normal. In contrast, many of the subsequent biopsy samples from both untrained and resistance-trained men and women contained evidence of damage. Ultrastructural analysis confirmed that degenerative-regenerative processes were occurring in both groups. However, training subjects had a four-fold greater number of damaged fibers than nontraining subjects (8.53% vs 2.08%). In addition, only biopsy samples from training individuals contained fibers with internal disorganization (e.g., Z-line streaming, myofibrillar disruption). Calpain II levels in the biopsy samples and serum creatine kinase activity were not significantly affected supporting the light and electron microscopic observations that most of the damaged fibers were normal in appearance except for their small diameter. In summary, focal damage induced by the biopsy procedure is not completely repaired after 2 weeks and could affect the results, particularly cross-sectional area measurements. Moreover, resistance training appears to cause additional damage to the muscle and may delay repair of the biopsied region.

Strength and skeletal muscle adaptations in heavy-resistance-trained women after detraining and retraining.

Six women who had participated in a previous 20-wk strength training study for the lower limb detrained for 30-32 wk and subsequently retrained for 6 wk. Seven untrained women also participated in the 6-wk "retraining" phase. In addition, four women from each group volunteered to continue training an additional 7 wk. The initial 20-wk training program caused an increase in maximal dynamic strength, hypertrophy of all three major fiber types, and a decrease in the percentage of type IIb fibers. Detraining had relatively little effect on fiber cross-sectional area but resulted in an increased percentage of type IIb fibers with a concomitant decrease in IIa fibers. Maximal dynamic strength decreased but not to pretraining levels. Retraining for 6 wk resulted in significant increases in the cross-sectional areas of both fast fiber types (IIa and IIab + IIb) compared with detraining values and a decrease in the percentage of type IIb fibers. The 7-wk extension accentuated these trends such that cross-sectional areas continued to increase (nonsignificant) and no IIb fibers could be found. Similar results were found for the nonpreviously trained women. These data suggest that rapid muscular adaptations occur as a result of strength training in previously trained as well as non-previously trained women. Some adaptations (fiber area and maximal dynamic strength) may be retained for long periods during detraining and may contribute to a rapid return to "competitive" form.

Serum creatine kinase activity and its changes after a muscle biopsy.

The significance of the absolute elevations of serum creatine kinase (CK) levels after intense exercise and injuries was studied by measuring CK activities from seven healthy active males during a 2-week period, with a muscle biopsy taken between the first and second week. Most of the subjects (three lifters and two runners) carried on their normal exercise activities, while two lifters stopped training during the 2 weeks. The weight of the biopsy, number of fibres, percentage of fibre types, and cross-sectional areas of the muscle fibres were measured. The CK levels of the non-active subjects and runners remained consistently low during the control week, whereas those of the lifters were usually 500% greater than those of the other two groups, and fluctuated with the intensity of their workouts. A muscle biopsy, having a mean weight of 71.3 mg and containing 1800 fibres, increased the CK values by approximately 100 units litre-1 (U l-1) in most of the subjects. One runner injured his right hamstring muscles 2 days prior to the biopsy, and his CK values rose from 50 to 4400 U l-1. The increases in CK after the biopsy were not related to fibre type, activity, weight of the biopsy, or number or size of fibres removed. These results indicate that: (1) CK values are consistently lower in normal subjects and runners than in lifters. (2) Weight training results in chronic elevations of CK. (3) Compared to a muscle biopsy, muscular injury dramatically increases CK levels. (4) Elevation of serum CK is observed as early as 1 h after an intense weight-lifting session.(ABSTRACT TRUNCATED AT 250 WORDS)

Lipid depletion and repletion in skeletal muscle following a marathon.

Intramuscular lipid content was investigated in muscle biopsies from 10 well-trained endurance athletes before, immediately after, and 1, 3, 5, and 7 days after a marathon. Diets were controlled throughout the entire period of the study. Triglyceride content was ultrastructurally determined by the use of stereological methods. The volume percent lipid significantly decreased after the marathon and was lowest at 3 days post-marathon, rising slightly but still 35% lower than the pre-marathon value by 7 days post-marathon. Glycogen granules were abundant and tightly packed in the pre-marathon biopsies, scarce immediately post-marathon, and abundant, but less tightly packed, 7 days post-marathon. Post-marathon fluctuations in the volume percentages of mitochondria indicated possible fluid shifts within the muscle fibers: dehydration immediately post-marathon followed by rehydration with possible edema. Assuming the content of mitochondria remained constant throughout the recovery period, the ratio of the volume percentage of lipid to the volume percentage of mitochondria indicated that lipid content may have reached pre-marathon levels after 7 days post-marathon.