Fibre types in human abdominal muscles

Histochemical muscle fibre composition was studied in biopsies from the four different muscles of the abdominal wall (rectus abdominis, RA, obliquus externus, OE, obliquus internus, OI, and transversus abdominis, Tr) in 13 normal human subjects (9 females and 4 males, age 24–55 years) undergoing gall-bladder surgery. Muscle fibres were classified as Type I, II A, II B or II C on the basis of their myofibrillar ATPases' pH lability. There were large inter-individual variations in fibre composition, whereas, in general, the differences between the different muscles were minor or non-existent. Mean fibre distribution ranges were 55–58% I, 15–23% II A, 21–28% II B, and 0–1% II C fibres. The least fibre diameters were similar for all types and muscles (range of means 50–54 μm) except for Tr in which the Type II fibres were smaller (mean 45 μm). There was a high correlation in the size of Type I vs. II fibres and Type II A vs. II B fibres in all layers. The oxidative potential (NADH-diaphorase staining intensity) appeared high in Type I fibres and low in Type II fibres, irrespective of subgroups. Thus, based on histochemical fibre composition, the different abdominal muscles appear to have a similar functional capacity. However, functional differences between individuals were indicated by the large inter-individual variation in muscle fibre distribution.     

Fibre types in human lumbar back muscles

The distribution of histochemically identified muscle fibre types was studied in biopsy samples from the two main muscles in the lumbar region of the human erector spinae, the multifidus and the longissimus, in 16 healthy subjects (nine males and seven females, age 20–30 years). Muscle fibres were classified as types I, IIA, IIB or IIC on the basis of the pH lability of their myofibrillar ATPases. There were no differences between the multifidus and the longissimus muscles in the relative occurrence of type I (62 vs. 57%), type IIA (20 vs. 22%) or type IIB fibres (18 vs. 22%), or in the absolute size of fibres (range of mean least diameters 58–66 μm). The oxidative potential (NADH-diaphorase staining intensity) was high in type I and low in type II fibres, irrespective of subgroups, in both muscles. In the females, the type I fibres occupied a relatively larger area (70–75 vs. 54–58% for the males) although the relative number of type I fibres was the same in both sexes. This was due to smaller type II fibres in the females resulting in higher type I/type II area ratios (1.70–1.90 vs. 0.88–0.92 for males). This suggests a difference in functional capacity of lumbar back muscles between males and females. On the other hand, the similarity in histochemical fibre-type distribution between the multifidus and the longissimus muscles does not give support for a functional differentiation between these two anatomically different parts of the lumbar erector spinae in man.     

Fibre types, capillary supply and enzyme activities in human intercostal muscles

The relative occurrence of slow twitch (ST) and fast twitch (FTa and FTb) fibres, fibre size, capillary supply, and glycolytic and oxidative enzyme activities in external (EXT) and internal (INT) intercostal muscles was determined on biopsies obtained during thoracotomy from the mid-axillary line in six patients with normal lung function. EXT and INT showed a similar occurrence of ST fibres. Both FTa and FTb fibres were seen in EXT while only FTa fibres were seen in INT. The size of the three fibre types in EXT was similar. In INT the fibre size was larger for FTa than for ST (P less than 0.05). INT appeared to have a larger number of capillaries per fibre than EXT (P less than 0.05). A positive correlation was demonstrated between variables measured during spirometry and fibre size in INT only. No difference in glycolytic and oxidative enzyme activities between EXT and INT was observed. The histochemical characteristics suggest that, in the mid-axillary line, INT are more used than EXT, though this conclusion did not appear to be confirmed by the enzyme activities determined.     

Fibre types inLimulus telson muscles: morphology and histochemistry

Summary Using a variety of techniques, we have demonstrated the presence of at least two fibre types inLimulus median telson levator muscle. By light and electron microscopy, large (21 56 m² mean cross-sectional area) fibres have A-bands of 4.1 m, one-half I bands of 2.15 m and Z lines 0.5 m in width. Few mitochondria are found in these fibres, which comprise 54% of those present in a given microscope field and which occupy 82% of the total cross-sectional area. Small fibres (484 m² mean cross-sectional area) have A bands of 6.3 m, one-half I bands of 3.1 m and Z lines between 0.5 and 1.0 m in width and are rich in mitochondria. Although small fibres comprise nearly one-half (46%) of the fibres in a field, they occupy only 18% of the total cross-sectional area.Histochemical staining for alkaline-stable myofibrillar ATPase activity and mitochondrial reduced -nicotinamide adenine nucleotide (-NADH) tetrazolium reductase activity confirms the presence of two fibre types. The large fibres react positively for the myofibrillar ATPase activity and negatively for the mitochondrial enzyme activity. The reverse is seen with the small fibres. Some fibres of intermediate size, having intermediate staining characteristics, were also observed. Native gel electrophoresis of both myofibrillar and purified myosin preparations supports the observed differences in myofibrillar ATPase activity in that two myosin isozymes are resolved on pyrophosphate gels. Although the thick filaments isolated from unstimulated small fibres are longer (>6.0 m) than those isolated from unstimulated large fibres (4.26 m), all have a similar appearance with respect to the arrangement of myosin heads on their surfaces, and similar diameters. The implications of the observed heterogeneity of fibre types is discussed with reference to previously reported phenomena inLimulus telson muscle, including changes in length of thick filaments on fibre stimulation and the shape of the length-tension curve obtained from fibre bundles.     

Fibre composition of human intrinsic tongue muscles

The muscle fibre composition of three human intrinsic tongue muscles, the longitudinalis, verticalis and transversus, was investigated in four anterior to posterior regions of the tongue using morphological and enzyme- and immunohistochemical techniques. All three muscles typically contained type I, IIA and IM/IIC fibres. Type I fibres expressed slow myosin heavy chain (MyHC), type II fibres fast MyHC, mainly fast A MyHC, whereas type IM/IIC coexpressed slow and fast MyHCs. Type II fibres were in the majority (60%), but regional differences in proportion and diameter of fibre types were obvious. The anterior region of the tongue contained a predominance of relatively small type II fibres (71%), in contrast to the posterior region which instead showed a majority of larger type I and type IM/IIC fibres (66%). In general, the fibre diameter was larger in the posterior region. This muscle fibre composition of the tongue differs from those of limb, orofacial and masticatory muscles, probably reflecting genotypic as well as phenotypic functional specialization in oral function. The predominance of type II fibres and the regional differences in fibre composition, together with intricate muscle structure, suggest generally fast and flexible actions in positioning and shaping the tongue, during vital tasks such as mastication, swallowing, respiration and speech.     

Stretch reflexes in human abdominal muscles

Homonymous and heteronymous reflex connections of the abdominal muscles were investigated by the application of a tap to the muscle belly and observation of surface electromyographic responses. Reflex responses of the following abdominal muscles were investigated both ipsilateral and contralateral to the tap: rectus abdominis (RA), external oblique (EO) and internal oblique (IO). Reflexes were evoked in each of the homonymous muscles with latencies and estimated conduction velocities compatible with being evoked by Ia muscle afferents and having a monosynaptic component. Short latency heteronymous excitatory reflex connections were also observed in muscles on both ipsilateral and contralateral sides in response to the same stimulus. The latencies of the crossed responses were only marginally longer than responses evoked in the respective ipsilateral muscle. Moreover, the reflexes evoked in the IO muscle from ipsilateral and contralateral IO muscle afferents were of comparable amplitude, as were those reflexes evoked in ipsilateral and contralateral EO and RA muscles when tapping IO. These similarities in the reflex characteristics on the ipsilateral and contralateral sides suggest that abdominal muscle afferents activate similar pathways to muscles on both sides of the body. It follows that if the homonymous stretch reflex of abdominal muscles have a monosynaptic component, then a similar monosynaptic pathway activates synergistic motoneurones, not only ipsilaterally but also contralaterally.     

Fibre types in extraocular muscles: a new myosin isoform in the fast fibres

Summary We report on the existence of a myosin heavy chain (MHC) isoform with unique structural properties in extraocular (EO) muscles. Differences in MHC composition are apparent using a polyclonal antibody prepared against myosin isolated from bovine EO muscle myosin. In enzyme immunoassays and western blotting experiments, this anti-EO myosin antibody reacted specifically with the heavy chains of EO muscle myosin and not with the heavy chains of other myosins. The distribution of this new MHC isoform in the globe rotating muscles from different mammalian species was analysed using a panel of specific anti-myosin antibodies and comparing the histochemical myosin ATPase profile of muscle fibres with their isomyosin content. Most fibres which display a type 2 ATPase reaction pattern were selectively labelled by anti-EO antibodies. A few type 2 fibres were found to react with both anti-EO and anti-2A myosin antibodies and others, located almost exclusively in the orbital layers, reacted with anti-foetals as well as anti-EO antibodies.The presence of a distinct form of myosin in EO muscle fibres is probably related to the particular functional characteristics of these muscles, which are known to be exceptionally fast-contracting but display a very low tension output.     

Relationship between isometric endurance and fibre types in human leg muscles.

Relationship between isometric endurance performance at 50% of maximal voluntary isometric contraction (MCV) and skeletal muscle fibre composition has been elucidated in 19 physical education students. This was found to be linear and the equation corresponded to: y=9.35 + 1.093x; r=0.70 (endurance time expressed in seconds and fibre composition as percent slow twitch muscle fibres (ST) of the vastus lateralis muscle). As it is assumed from previous studies that similar isometric tensions preferentially recruit fast twitch muscle fibres (FT) and that the muscle at the point of exhaustion exhibits maximal values for lactate accumulation, it is suggested that lactate formed in FT fibres is released and stored in nonrecruited ST fibres. The ability to sustain similar isometric tension would then be depending on how large the fraction of ST fibres is that can serve as a lactaterecipient for lactate producing FT fibres.     

Immunohistochemical study of fiber types in human extraocular muscles

Fiber types in human extraocular muscle (h-EOM) were examined immunohistochemically with antibodies against slow tonic (anti-ALD) and slow twitch (anti-SOL) myosins. Four types of muscle fiber in h-EOM were distinguishable according to their reactivities with these antibodies. Groups 1 and 2 fibers reacted with both antibodies, group 1 fibers showing stronger reactivity than group 2 fibers with anti-ALD. Group 3 fibers reacted only with anti-SOL. Group 4 fibers did not react with either antibody. The latter were the most common, and were the main fibers in both the peripheral (outer orbital) and central zones of h-EOM. The next most common were group 1 fibers, which were located mainly in the peripheral layer. Group 2 fibers were less common, but were the second most common type in the central layer. Group 3 fibers were only minor constituents. Multiple innervations were observed in some fibers of groups 1 and 2, and group 1 fibers were suggested to be slow tonic myofibers in h-EOM. These specific immunohistochemical and physiological features of h-EOM seem to be the basis of the low morbidity seen in the usual types of muscular dystrophy.     

Preliminary observations on the microarchitecture of the human abdominal muscles

Precise knowledge of muscle architecture and innervation patterns is essential for the development of accurate clinical and biomechanical models. Although the gross anatomy of the human abdominal muscles has been investigated, the finer details of their microanatomy are not well described. Fascicles were systematically sampled from each of the human abdominal muscles, and small fiber bundles from selected fascicles stained with acetylcholinesterase to determine the location of motor endplate bands, myomyonal junctions, and myotendinous junctions. Statistical analysis was used to ascertain the association between fascicular length and number of endplate bands. The number of endplate bands along a fascicle was variable between different portions of each muscle, but was strongly correlated with fascicular length (r = 0.814). In fascicles less than 50 millimeters (mm) in length, only a single endplate band was generally present, while multiple endplate bands (usually two or three) were found in fascicles longer than 50 mm. The presence of myomyonal junctions throughout the longer (>50 mm) fascicles verified that they were composed of short, intrafascicularly terminating fibers, while shorter fascicles comprised fibers spanning the entire fascicular length. This preliminary study provides evidence that multiple endplate bands are contained in some regions of the abdominal muscles, an arrangement that differs from most human appendicular muscles. It is not clear whether the variations in the described fine architectural features reflect regional differences in muscle function.     

Comparison of human motor cortical projections to abdominal muscles and intrinsic muscles of the hand

Summary Percutaneous electrical stimulation of the motor cortex was used to activate rapidly conducting corticofugal pathways to human abdominal muscles. Following cortical stimulation the response latencies for the abdominal muscles were similar to those for limb muscles which are a similar distance from the motor cortex. Cortically evoked responses recorded from the abdominal muscles had the same latency and similar amplitude during several voluntary tasks including expiration, expulsive manoeuvres and trunk flexion. Responses could also be evoked when the chemical drive to breathe was increased by rebreathing. In addition, the properties of the cortical projection to muscles of the abdominal wall were directly compared with those of the projection to the intrinsic muscles of the hand. The latencies of responses in abdominal muscles and intrinsic muscles of the hand were measured during static contractions over a range of strengths in the same subjects (0–100% maximal voluntary contraction, MVC). For both muscle groups, cortically evoked muscle responses of minimal latency occurred when background contractions reached 10–20% MVC with responses of maximal amplitude at 60% MVC. The variability in latency of fifty consecutive responses were similar for the two muscle groups. Furthermore, post-stimulus time histograms for 4 rectus abdominis motoneurones revealed a brief initial excitatory peak of 1.15ms duration (range 0.96–1.34ms) following cortical stimulation. The characteristics of this peak are the same as reported for motoneurones of intrinsic hand muscles. These findings demonstrate a powerful rapidly conducting pathway from the motor cortex to the human abdominal muscles. This pathway has many of the same properties as the monosynaptic corticospinal projection to the distal muscles of the upper limb.     

Fiber types in mammalian skeletal muscles

Mammalian skeletal muscle comprises different fiber types, whose identity is first established during embryonic development by intrinsic myogenic control mechanisms and is later modulated by neural and hormonal factors. The relative proportion of the different fiber types varies strikingly between species, and in humans shows significant variability between individuals. Myosin heavy chain isoforms, whose complete inventory and expression pattern are now available, provide a useful marker for fiber types, both for the four major forms present in trunk and limb muscles and the minor forms present in head and neck muscles. However, muscle fiber diversity involves all functional muscle cell compartments, including membrane excitation, excitation-contraction coupling, contractile machinery, cytoskeleton scaffold, and energy supply systems. Variations within each compartment are limited by the need of matching fiber type properties between different compartments. Nerve activity is a major control mechanism of the fiber type profile, and multiple signaling pathways are implicated in activity-dependent changes of muscle fibers. The characterization of these pathways is raising increasing interest in clinical medicine, given the potentially beneficial effects of muscle fiber type switching in the prevention and treatment of metabolic diseases.     

Activation of two types of fibres in rat extraocular muscles.

1. The contractile responses of the inferior rectus, one of the extraocular muscles of the rat, to a depolarization induced by an elevation of the potassium concentration in the external medium ([K]O) have been studied 'in vitro'. 2. The elevation of [K]O to 20 and 30 mM-K produced contractures that consisted of a sustained or tonic tension. When [K]O was increased to 50 mM or more a well-defined transient or phasic tension appeared before the tonic response. The increment of [K]O above 50 mM enhanced the phasic component and depressed the tonic tension. The maximal tonic tension, usually evoked by 50 mM-K, is about 50% of the tetanic tension, shows a gradual decline with time and lasts for hours. Control experiments performed in diaphragm showed that this muscle only responds with phasic tensions. 3. The difference in the repriming of the phasic and tonic responses when tensions were induced with salines containing low or normal [Cl] suggests that the muscle fibres responsible for the tonic tension are poorly permeable to Cl-. 4. The amplitude of the tonic tension was reduced by Ca deprivation and by an elevation of [Ca] in the saline to 10 mM. 5. It is concluded that in rat extraocular muscles, an increase in [K]O activates two types of muscle fibres: singly and multiply innervated. These appear to be functionally equivalent to the twitch and slow fibres of amphibian and avian muscle and would give rise to the phasic and tonic components of the contracture, respectively.     

Load compensation in human masseter muscles.

The reaction of masseter and digastric muscles to changes in the load applied during a movement was studied in human volunteers. 2. Rapid stretch of the jaw-closing muscles evoked a monosynaptic jaw-jerk reflex response in the masseteric electromyogram (e.m.g.) after a delay of 6-8 msec. 3. The averaged integrated e.m.g. activity of the masseter was increased 5-10 msec after the rapid application of a load during voluntary closure of the jaw. It was not necessary to stretch the muscle to obtain this apparently monosynaptic response. 4. A compensatory increase in the velocity of movement followed 6-10 msec after the e.m.g. peak. 5. Unloading the masseter muscle during contraction was followed by a fall in e.m.g. activity, beginning after a latency of 6-5--1 msec, with later phases of depression beginning 27--36 and 60--70 msec after unloading. 6. The spindle-poor digastric muscles were not monosynaptically activated during loading, but responded with a latency of 24--34 msec. 7. It is concluded that monosynaptic e.m.g. changes are a prominent feature of the load compensation system in human jaw-closing muscles but not in jaw-opening muscles.     

Fibre types in skeletal muscle: a personal account

Muscle performance is in part dictated by muscle fibre composition and a precise understanding of the genetic and acquired factors that determine the fibre type profile is important in sport science, but is also relevant to neuromuscular diseases and to metabolic diseases, such as type 2 diabetes. The dissection of the signalling pathways that determine or modulate the muscle fibre phenotype has thus potential clinical significance. In this brief review, I examine the evolution of the notion of muscle fibre types, discuss some aspects related to species differences, point at problems in the interpretation of transgenic and knockout models and show how in vivo transfection can be used to identify regulatory factors involved in fibre type diversification, focusing on the calcineurin-nuclear factor of activated T cells (NFAT) pathway.     

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.