Mammalian skeletal muscle is an extremely heterogeneous tissue, composed of a large variety of fiber types. These fibers, however, are not fixed units but represent highly versatile entities capable of responding to altered functional demands and a variety of signals by changing their phenotypic profiles. This adaptive responsiveness is the basis of fiber type transitions. The fiber population of a given muscle is in a dynamic state, constantly adjusting to the current conditions. The full range of adaptive ability spans fast to slow characteristics. However, it is now clear that fiber type transitions do not proceed in immediate jumps from one extreme to the other, but occur in a graded and orderly sequential manner. At the molecular level, the best examples of these stepwise transitions are myofibrillar protein isoform exchanges. For the myosin heavy chain, this entails a sequence going from the fastest (MHCIIb) to the slowest (MHCI) isoform, and vice-versa. Depending on the basal protein isoform profile and hence the position within the fast-slow spectrum, the adaptive ranges of different fibers vary. A simple transition scheme has emerged from the multitude of data collected on fiber type conversions under a variety of conditions.

Mammalian skeletal muscle fiber type transitions

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 the vastus lateralis muscle of young men and women.

Although many training variables contribute to the performance, cellular and molecular adaptations to resistance exercise, relative intensity (% 1 repetition maximum [%1RM]) appears to be an important factor. This review summarises and analyses data from numerous resistance exercise training studies that have monitored percentage fibre type, fibre type cross-sectional areas, percentage cross-sectional areas, and myosin heavy chain (MHC) isoform expression. In general, relative intensity appears to account for 18-35% of the variance for the hypertrophy response to resistance exercise. On the other hand, fibre type and MHC transitions were not related to the relative intensity used for training. When competitive lifters were compared, those typically utilising the heaviest loads (> or =90% 1RM), that is weightlifters and powerlifters, exhibited a preferential hypertrophy of type II fibres when compared with body builders who appear to equally hypertrophy both type I and type II fibres. These data suggest that maximal hypertrophy occurs with loads from 80-95% 1RM.

The Role of Resistance Exercise Intensity on Muscle Fibre Adaptations

1. Age-associated loss of skeletal muscle mass and strength can partly be counteracted by resistance training, causing a net synthesis of muscular proteins. Protein synthesis is influenced synergistically by postexercise amino acid supplementation, but the importance of the timing of protein intake remains unresolved. 2. The study investigated the importance of immediate (P0) or delayed (P2) intake of an oral protein supplement upon muscle hypertrophy and strength over a period of resistance training in elderly males. 3. Thirteen men (age, 74 ± 1 years; body mass index (BMI), 25 ± 1 kg m_2 (means ± S.E.M.)) completed a 12 week resistance training programme (3 times per week) receiving oral protein in liquid form (10 g protein, 7 g carbohydrate, 3 g fat) immediately after (P0) or 2 h after (P2) each training session. Muscle hypertrophy was evaluated by magnetic resonance imaging (MRI) and from muscle biopsies and muscle strength was determined using dynamic and isokinetic strength measurements. Body composition was determined from dual-energy X-ray absorptiometry (DEXA) and food records were obtained over 4 days. The plasma insulin response to protein supplementation was also determined. 4. In response to training, the cross-sectional area of m. quadriceps femoris (54.6 ± 0.5 to 58.3 ± 0.5 cm2 ) and mean fibre area (4047 ± 320 to 5019 ± 615 µm 2 ) increased in the P0 group, whereas no significant increase was observed in P2. For P0 both dynamic and isokinetic strength increased, by 46 and 15 %, respectively ( P< 0.05), whereas P2 only improved in dynamic strength, by 36 % ( P< 0.05). No differences in glucose or insulin response were observed between protein intake at 0 and 2 h postexercise. 5. We conclude that early intake of an oral protein supplement after resistance training is important for the development of hypertrophy in skeletal muscle of elderly men in response to resistance training.

https://physoc.onlinelibrary.wiley.com/doi/pdf/10.1111/j.1469-7793.2001.00301.x

Timing of postexercise protein intake is important for muscle hypertrophy with resistance training in elderly humans

Contractile and energetic properties of human skeletal muscle have been studied for many years in vivo in the body. It has been, however, difficult to identify the specific role of muscle fibres in modulating muscle performance. Recently it has become possible to dissect short segments of single human muscle fibres from biopsy samples and make them work in nearly physiologic conditions in vitro. At the same time, the development of molecular biology has provided a wealth of information on muscle proteins and their genes and new techniques have allowed analysis of the protein isoform composition of the same fibre segments used for functional studies. In this way the histological identification of three main human muscle fibre types (I, IIA and IIX, previously called IIB) has been followed by a precise description of molecular composition and functional and biochemical properties. It has become apparent that the expression of different protein isoforms and therefore the existence of distinct muscle fibre phenotypes is one of the main determinants of the muscle performance in vivo. The present review will first describe the mechanisms through which molecular diversity is generated and how fibre types can be identified on the basis of structural and functional characteristics. Then the molecular and functional diversity will be examined with regard to (1) the myofibrillar apparatus; (2) the sarcolemma and the sarcoplasmic reticulum; and (3) the metabolic systems devoted to producing ATP. The last section of the review will discuss the advantage that fibre diversity can offer in optimizing muscle contractile performance.

Human skeletal muscle fibres: molecular and functional diversity

Histochemical methods are routinely used to delineate skeletal muscle fiber types. In the present investigation, this qualitative determination of fiber type composition was compared to the electrophoretically determined myosin heavy chain (MHC) content from a large number of human muscle biopsy samples. Biopsies were taken from the vastus lateralis muscle at the beginning and every 2 weeks during 8 weeks of high-intensity resistance training from men (n = 13) and woman (n = 8). Muscle was also extracted from nontraining men (n = 7) and women (n = 5) at the same periods. Six muscle fiber types (I, IC, IIAC, IIA, IIAB, and IIB) were determined using basic myofibrillar adenosine triphosphatase histochemistry. Cross-sectional areas were determined for the three major fiber types (I, IIA, and IIB) and used to calculate the percentage area of these types. Electrophoretic techniques were used to separate and quantify the percentage MHC content in these same biopsy samples, and these data were then used to compare with the percentage fiber type area. Correlation analyses suggest a relationship between the histochemically assessed percentage fiber type area and the electrophoretically assessed MHC content in human limb musculature. However, because of possible histochemical misclassification of some fibers (especially in trained muscle) both techniques may be essential in yielding important information about fiber type composition and possible fiber type transformations.

Correlation between percentage fiber type area and myosin heavy chain content in 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).

C. A. Allemeier

Papers

Correlation between percentage fiber type area and myosin heavy chain content in human skeletal muscle.

Effects of sprint cycle training on human skeletal muscle