Correlation of myosin isoforms with anatomical divisions in equine musculus biceps brachii.

The biceps brachii of horses is subdivided into a lateral and medial head. Electrophoresis of samples from the lateral head revealed three slow-migrating native myosin isoforms, including one that does not correspond to slow myosin isoforms described for other mammalian muscles. In contrast, the medial head contained a single slow isoform. Both the lateral and medial heads contained three fast-migrating isoforms corresponding with the FM-2, FM-3 and FM-4 isoforms reported for other mammalian fast-twitch muscle fibers. Electrophoresis of myosin heavy chains (MHCs) revealed only two MHC bands, one fast-migrating band that comigrates with rat type I MHC and a second slower-migrating band that comigrates with rat type IIa MHC. Quantitation of the histochemical data is correlated with densitometric analysis of MHCs in the medial and lateral heads of biceps brachii and is consistent with previously hypothesized functional specializations of this muscle.     

Elbow extensor muscles of the horse: postural and dynamic implications.

Based on histochemical and immunohistochemical evidence, horse elbow extensor muscles are composed of two morphologically distinct muscle groups. The long and lateral heads of the triceps brachii are large, predominantly type II (presumed fast) muscles. The long and lateral heads of the triceps together account for 96% of the weight of the elbow extensors (long head of triceps is 81%). The long and lateral heads contain three histochemical fiber types: types I, IIa and IIb. Type I muscle fibers account for approximately 18 and 27% of the fibers in the long and lateral heads of the triceps, respectively. In the lateral head, type IIa and IIb fibers account equally for the remaining 70%, while in the long head of the triceps type IIb fibers predominate (50%) over type IIa fibers (32%). In contrast, the much smaller medial head of the triceps (2% of triceps mass) and the anconeus (2% of mass) contain almost exclusively type I muscle fibers. It is hypothesized that the medial head and anconeus, with their slow fibers, contribute to the postural maintenance of the forelimb by preventing flexion at the elbow joint during passive stance. The larger long and lateral heads, with their generally fast fiber populations, are most likely important during dynamic activity.     

Functional morphology of the endomysium in series fibered muscles.

Many skeletal muscles, including the feline biceps femoris, are composed of short, tapered myofibers arranged in an overlapping longitudinal series. The endomysium of such muscles transfers tension between overlapping myofibers, and is thus an elastic element in series with them. The endomysium of the cat biceps femoris contains curvilinear collagen fibrils in an approximately isotropic (random) array. The collagen fibrils undergo only a modest reorientation as the myofibers shorten or lengthen within the physiological range. A geometrical model predicts no change in the thickness of the endomysium on changing muscle fiber length and quantifies the expected collagen fibril reorientation in the endomysium as a function of muscle extension. It is also demonstrated that a high proportion of the collagen fibrils will be curvilinear at all sarcomere lengths. The organization of endomysial collagen is appropriate for the transfer of loads between myofibers by means of shear.     

Interfiber tension transmission in series-fibered muscles of the cat hindlimb

Several muscles of the cat hindlimb, including biceps femoris and tenuissimus, are composed of short, in-series muscle fibers with tapered intrafascicular terminations. Tension generation and transmission within such muscles requires that active fibers should be mechanically coupled in series via myomyous junctions, specialized connective tissue attachments, or the endomysium. This report establishes that the tapered fibers of the cat biceps femoris and tenuissimus muscles have insignificant numbers of either myomyous or specialized connective tissue junctions. Tension appears to be transmitted in a distributed manner across the plasmalemma of the tapered (and probably the non-tapered) portions of the fibers to the connective tissue of the endomysium, which is therefore an essential series elastic element in these muscles. Subplasmalemmal dense plaques were identified and may play a role in transmembrane force transmission. In addition to the endomysium, passive muscle fibers may also serve to transmit tension between active fibers, and therefore should also be considered to be series elastic elements.     

Four forearm flexor muscles of the horse, Equus caballus: anatomy and histochemistry.

Two of the forearm flexors of the horse, the superficial and deep digital flexor muscles, are critical to support the digital and fetlock joints, exhibit differing insertions, and are passively supported by the proximal and distal check ligaments, respectively. These two muscles differ in histochemical composition and architecture. The differences are correlated with the different stress levels transmitted through their tendons, and the different frequencies of clinical breakdown that have been reported. Both muscles contain type I and type IIa fibers. A few type IIb fibers occurred in the deep digital flexor. The superficial digital flexor contained approximately 56% type I fibers, extremely short muscle fibers, and extensive connective tissue investment. In contrast, the deep digital flexor had three muscle heads: ulnar, radial, and "long" and "short" regions of the humeral head. The "long" and "short" regions of the humeral head contained 33% and 44% type I fibers, respectively, fiber lengths three to four times as long as those in the superficial digital flexor, and relatively less connective tissue investment. Flexor carpi radialis and flexor carpi ulnaris compared most closely with the humeral head of the deep digital flexor. These data suggest a correlation of the unique architecture of superficial digital flexor with its proposed elastic storage properties during locomotion in horses, and an explanation for the frequent breakdown of the superficial digital flexor in athletic horses.     

A morphologic and histochemical study of the mesentery in the guinea pig

The guinea pig mesentery is a uniform, continuous, thin (18 micron) sheet of connective tissue covered by a single layer of flattened mesothelial cells on both surfaces. Tight and gap junctions provide for cell-to-cell adhesion among mesothelial cells. These cells possess numerous micropinocytotic vesicles; a conspicuous basal lamina separates the mesothelium from the underlying connective tissue. Most of the material found between the two serous coverings consisted of a three-dimensional meshwork of abundant collagenous fibers intermingled with a sparse net of very thin (0.4 micron) elastic fibers. Two distinct populations of collagen fibrils are segregated into different compartments of the mesentery. One population is formed of thick (56 nm) fibrils which associate to form closely packed fibers. The second population, composed of loosely arranged thin (38 nm) fibrils which do not become assembled into fibers, is found underlying the basal lamina that separates the mesothelium from the connective tissue. These observations strongly suggest that the mesentery contains both collagens type I and type III. The guinea pig mesentery contains 6.8 mg of sulfated glycosaminoglycans/g dry weight. Most of these glycosaminoglycans (78%) were identified as dermatan sulfate, whilst the rest (22%) corresponded to heparan sulfate.     

Changes in the activity of succinate dehydrogenase in muscle fibers of functionally different muscles in the rat after a reduction in exertion--hypodynamia

In 105 male rats of Wistar strain distal parts of one of the thoracic extremities are amputated with keeping intact the places where the brachial muscle is fixed. This does not restrict the volume of the brachium movements but essentially decreases their dynamic component (power loading). For 45 days dynamics of succinate-dehydrogenase (SDG) activity is being revealed in muscle fibers (MF) of functionally different muscles: m. brachialis, m. serratus ventralis and m. triceps brachii (the medial head). Average tendency of the process, changes in the distribution margins, asymmetry and kurtosis are taken into account. Under hypodynamia reconstruction of the MF has a wavy character with a gradually longer period of fluctuations. In all the muscles appear MF with a greater than in the control SDG activity. In the medial head of the m. triceps brachii the fibers with the lowest SDG activity disappear. The amount of MF with the lowest activity decreases, while those with the higher--increases, the process being more pronounced in the m. triceps brachii. The amount of MF with middle activity of the enzyme remains nearly unchanged. MF with different initial enzymatic activity do not change simultaneously. The degree of the changes in the fibers and the power leading are connected with each other, the fibers with the low initial SDG activity including into the reaction at a sharper decrease of the latter. The changes in quantitative ratio of MF with different SDG activity are not the same in every muscle studied.     

Composition of myofiber types in the vocalis muscles involved in rapid closure of the glottis in Japanese macaques

Myofiber types of the medial thyroarytenoid (vocalis) muscle, lateral thyroarytenoid muscle, and cricothyroid muscle of the Japanese macaque were examined with enzyme-histochemical methods. For comparison, the semitendinosus muscle of the Japanese macaque and the thyroarytenoid (vocalis) muscle of cattle, sheep, and pig were examined with the same methods. The vocalis muscle of the Japanese macaque was composed exclusively of fast-twitch/oxidative/glycolytic (FOG) myofibers; it differed from the lateral thyroarytenoid, cricothyroid, and semitendinosus muscles of the Japanese macaque and from the vocalis muscles of the other animals, which consisted of slow-twitch/oxidative, FOG-, and fast-twitch/glycolytic myofibers or type IIC myofibers. The histochemical properties of the vocalis muscle of the Japanese macaque show that the vocalis muscle has a capacity to close the glottis rapidly and completely.     

A 3D model of muscle reveals the causes of nonuniform strains in the biceps brachii

Biomechanical models generally assume that muscle fascicles shorten uniformly. However, dynamic magnetic resonance (MR) images of the biceps brachii have recently shown nonuniform shortening along some muscle fascicles during low-load elbow flexion (J. Appl. Physiol. 92 (2002) 2381). The purpose of this study was to uncover the features of the biceps brachii architecture and material properties that could lead to nonuniform shortening. We created a three-dimensional finite-element model of the biceps brachii and compared the tissue strains predicted by the model with experimentally measured tissue strains. The finite-element model predicted strains that were within one standard deviation of the experimentally measured strains. Analysis of the model revealed that the variation in fascicle lengths within the muscle and the curvature of the fascicles were the primary factors contributing to nonuniform strains. Continuum representations of muscle, combined with in vivo image data, are needed to deepen our understanding of how complex geometric arrangements of muscle fibers affect muscle contraction mechanics.     

Ultrastructure of sea urchin tube feet

Summary An analysis of the ultrastructure of the tube feet of three species of sea urchins (Strongylocentrotus franciscanus, Arbacia lixula and Echinus esculentus) revealed that the smooth muscle, although known to be cholinoceptive, receives no motor innervation.The muscle fibers are attached to a double layer of circular and longitudinal connective tissue which surrounds the muscle layer and contains numerous bundles of collagen fibers. On its outside, the connective tissue cylinder is invested by a basal lamina of the outer epithelium to which numerous nerve terminals are attached. These are part of a nerve plexus which surrounds the connective tissue cylinder. The plexus itself is an extension of a longitudinal nerve that extends the whole length of the tube foot. It is composed of axons, but nerve cell bodies and synapses are conspicuously lacking, suggesting that the axons and terminals derive from cells of the radial nerve. Processes of the epithelial cells penetrate the nerve plexus and attach to the basal lamina. There is no evidence that the epithelial cells function as sensory cells.On the basis of supporting evidence it is suggested that the transmitter released by the nerve terminals diffuses to the muscle cells over a distance of several microns and in doing so affects the mechanical properties of the connective tissue.Supported by the Sonderforschungsbereich 138 of the Deutsche Forschungsgemeinschaft     

Muscle fiber number in biceps brachii in bodybuilders and control subjects

Muscle fiber numbers were estimated in vivo in biceps brachii in 5 elite male bodybuilders, 7 intermediate caliber bodybuilders, and 13 age-matched controls. Mean fiber area and collagen volume density were calculated from needle biopsies and muscle cross-sectional area by computerized tomographic scanning. Contralateral measurements in a subsample of seven subjects indicated the method for estimation of fiber numbers to have adequate reliability. There was a wide interindividual range for fiber numbers in biceps (172,085-418,884), but despite large differences in muscle size both bodybuilder groups possessed the same number of muscle fibers as the group of untrained controls. Although there was a high correlation between average cross-sectional fiber area and total muscle cross-sectional area within each group, many of the subjects with the largest muscles also tended to have a large number of fibers. Since there were equally well-trained subjects with fewer than normal fiber numbers, we interpret this finding to be due to genetic endowment rather than to training-induced hyperplasia. The proportion of muscle comprised of connective and other noncontractile tissue was the same for all subjects (approximately 13%), thus indicating greater absolute amounts of connective tissue in the trained subjects. We conclude that in humans, heavy resistance training directed toward achieving maximum size in skeletal muscle does not result in an increase in fiber numbers.     

Histochemical characteristics of the muscle fibers of the biceps and triceps brachii muscles in human ontogeny

By means of morphometrical and histochemical methods for revealing myosin ATPase and SDG activity development of various types of muscle fibers (MF) has been studied in the postmortem material, using m. biceps and m. triceps brachii in human ontogenesis. The flexors and extensors have features in common in the dynamics of the MF maturation, and some distinctive peculiarities. The appearance of histochemical distinctions between the MF takes place on the 5th-6th months of the intrauterine development. Morphofunctional specialization begins with formation of tonic fibers. During the 1st-2nd years phasic fibers form. A relative amount of fast MF in both muscles increases at the age of 11-12 years. The dynamics of final specialization of the MF is connected with stages of sexual maturation. The first stage of the sexual maturation (about 14 years of age) is connected with decrease in the relative amount of the MF of glycolytic type of energy supply and corresponding increase in the number of oxidative type structures. From 15-17 years of age a final differentiation begins, it is connected with an intensive transversal growth of all the MF and distinguish of thick glycolytic MF. The m. biceps brachii has a relatively greater amount of oxidative fibers, and the m. triceps brachii, glycolytic ones. The transversal section area of the MF in the m. triceps brachii exceeds that of the m. biceps brachii, beginning from the 7th month of the intrauterine development up to 14 years of age. The investigation performed does not reveal any anticipating development either in the flexors or in the extensors. The differentiating processes in the m. biceps and m. triceps brachii occur nearly simultaneously.     

Distribution of innervation zone and muscle fiber conduction velocity in the biceps brachii muscle

The spatial distributions of muscle innervation zone (IZ) and muscle fiber conduction velocity (CV) were examined in nine healthy young male participants. High-density surface electromyography (EMG) was collected from the biceps brachii muscle when subjects performed isometric elbow flexions at 20% to 80% of the maximal voluntary contraction (MVC). A total of 9498 samples of IZs were identified and CVs were calculated using the Radon transform. The center and width of IZ sample distribution were compared within four different force levels and six medial to lateral electrode column positions using repeated measures ANOVA and multiple comparison tests. Significant shifts of IZ center were observed in the medial columns (Columns 5, 6, and 7) compared with the lateral columns (Columns 3 and 4) (p < 0.05). Similarly, significant differences in the IZ width were found in Column 7 and 8 compared to Column 3 (p < 0.05). In contrast, muscle CV was unaffected by column position. Instead, muscle CV was faster at 40% and 80% MVC compared to 20% MVC (p < 0.05). The findings of this study add further insights into the physiological properties of the biceps brachii muscle.     

The arrangement and function of octopus arm musculature and connective tissue

The morphology of the musculature and connective tissues of the arms of Octopus bimaculoides was analyzed with light microscopy. We also studied O. briareus and O. digueti, which possess relatively more elongate and less elongate arms, respectively. The morphology of the arms was found to be remarkably uniform among species. The arms consist of a densely packed three-dimensional arrangement of muscle fibers and connective tissue fibers surrounding a central axial nerve cord. Three primary muscle fiber orientations were observed: 1) transverse muscle fibers oriented in planes perpendicular to the long axis of the arm; 2) longitudinal muscle fibers oriented parallel to the long axis; and 3) oblique muscle fibers arranged in helixes around the arm. The proportion of the arm cross section occupied by each of these muscle fiber groups (relative to the total cross sectional area of the musculature) remains constant along the length of the arm, even though the arm tapers from base to tip. A thin circular muscle layer wraps the arm musculature on the aboral side only. Much of this musculature has its origin and insertion on several robust connective tissue sheets including a layer surrounding the axial nerve cord and crossed-fiber connective tissue sheets located on the oral and the aboral sides of the arm. An additional thin layer of connective tissue wraps the arm musculature laterally and also serves as a site of origin and insertion of some of the muscle fibers. The fibers of the oral and aboral crossed-fiber connective tissue sheets are arranged oblique to the long axis of the arm with the same fiber angle as the oblique muscle layers that originate and insert on the sheets. The oblique muscle layers and the crossed-fiber connective tissue sheets thus form composite right- and left-handed helical fiber arrays. Analysis of arm morphology from the standpoint of biomechanics suggests that the transverse musculature is responsible for elongation of the arms, the longitudinal musculature is responsible for shortening, and the oblique muscle layers and associated connective tissues create torsion. Arm bending may involve unilateral contraction of longitudinal muscle bundles in combination with resistance to arm diameter increase due to contraction of the transverse musculature or passive stiffness of the arm tissues. The arms may also be bent by a combination of decrease in diameter due to contraction of the transverse musculature and maintenance of constant length on one side of the arm by unilateral activity of longitudinal muscle bundles. An increase in flexural stiffness of the arm may be achieved by cocontraction of the transverse and longitudinal muscle. Torsional stiffness may be increased by simultaneous contraction of both the right- and left-handed oblique muscle layers.     

Metabolic capacity of individual muscle fibers from different anatomic locations.

We studied muscle fibers by quantitative biochemistry to determine whether metabolic capacity varied among fibers of a given type as a function of their anatomic location. Muscles were selected from both contiguous and diverse anatomic regions within the rats studied. The individual fibers, classified into myosin ATPase fiber types by histochemical means, were assessed for fiber diameters and analyzed for the activities of enzymes representing major energy pathways: malate dehydrogenase (MDH, oxidative), lactate dehydrogenase (LDH, glycolytic), and adenylokinase (AK, high-energy phosphate metabolism). We found that neither the average activities of each of the three enzymes nor the fiber diameters varied in Type I or Type IIa fibers selected from superficial to deep portions of the triceps surae of the hindlimb. However, the IIb fibers in the deep region of this muscle group had significantly greater oxidative capacity, less glycolytic capacity, and smaller diameters than the superficially situated IIb fibers. Type IIa fibers in lateral gastrocnemius, extensor digitorum longus, psoas, diaphragm, biceps brachii, superficial masseter, and superior rectus muscles were highly variable in both diameter and enzyme profiles, with a correlation between MDH activity and fiber diameter. Therefore, our results show that both intermuscular and intramuscular metabolic variations exist in muscle fibers of a given type.     

EMG and MMG activities of agonist and antagonist muscles in Parkinson’s disease patients during absolute submaximal load holding

The purpose of the study was (1) to assess changes in electromyographical (EMG) and mechanomyographical (MMG) signals of the biceps and triceps brachii muscles during absolute submaximal load holding in Parkinson’s disease patients tested during their medication “ON-phase” and in age-matched controls, and (2) to check whether mechanomyography can be useful in evaluation of neuromuscular system activity in Parkinson’s disease patients.The data analysis was performed on nine females with Parkinson’s disease and six healthy, age-matched females. The EMG and MMG signals were recorded from the short head of the biceps brachii (BB) and the lateral head of the triceps brachii (TB) muscles.It was concluded that compared to the controls, the Parkinson’s disease patients exhibited higher amplitude in the biceps brachii muscle and lower median frequency of the MMG signal in the both tested muscles. However, no differences in the EMG amplitude and an increase of the EMG median frequency in the triceps brachii muscle of the Parkinson’s disease group were observed. The MMG was not affected by physiological postural tremor and can depict differences between parkinsonians and controls, which may suggest that it is valuable tool for neuromuscular assessment for this condition.     

Microvascular architecture of the pampiniform plexus-testicular artery system in the rat: a scanning electron microscope study of corrosion casts

Because the architectural and biochemical properties of skeletal muscle dictate its force, velocity, and displacement properties, the major extensors (triceps brachii) and flexors (biceps brachii, brachialis, and brachioradialis) of the elbow in a primate (cynomolgus, monkey) were studied. Functional cross-sectional areas (CSA) were calculated from muscle mass, mean fiber length (normalized to a 2.20 microns sarcomere length), and angle of fiber pinnation measurements from each muscle. Fiber-type distributions were determined and used as a gross index of the biochemical capacities of the muscle. The extensor group had a shorter mean fiber length (31 vs. 47 mm), a larger CSA (13 vs. 8 cm2), and a higher overall percentage of slow-twitch fibers (47 vs. 26%). Consequently, the elbow extensors had a relatively greater potential for force production and force maintenance than the flexors. In contrast, the flexors were designed to optimize their length-velocity potentials; i.e., they had relatively long fibers and a higher fast-twitch fiber composition than the extensors. These morphologic differences between antagonistic muscle groups should be considered when evaluating the motor control mechanisms regulating reciprocal movements about the elbow.     

A new classification of the brachial extensors in Tarsius.

The muscular system of the tarsier was first described by Burmeister (1846), who noted that brachial extensors (triceps complex) have six heads. The first three heads, respectively, correspond to the long, lateral and medial heads of the triceps brachii muscle in man. The fourth head is the anconaeus and the fifth is the dorsoepitrochlearis. Schultz (1984) divided the sixth head into two different parts (preaxial and postaxial) from the viewpoint of nerve supply. The present study found that the whole sixth head is innervated by the ulnar nerve, and we propose that it is recognized as the proximal and distal heads of the (preaxial) epitrochleoanconaeus muscle. The proximal head may have developed specially in the tarsier in addition to the distal head observed in other prosimians. It is thought to support the extension of the elbow joint and contribute to the tarsier's effective locomotion.     

Histological features of endomysium,perimysium and epimysium in rat lateral pterygoid muscle

The distribution of the endomysium, perimysium, and epimysium and their constituent connective tissue fiber types in the mature rat lateral pterygoid muscle was examined with the light microscope. The endomysium and perimysium were relatively thin and consisted mainly of reticular fibers. The epimysium was thicker than the intramuscular sheaths and consisted of both collagen and reticular fibers; however, the thickness and constituent connective tissue fiber types of these sheaths varied regionally. Near the articular capsule and disc, the endomysium, perimysium, and epimysium were all thicker than in other regions of the muscle and consisted of collagen, reticular, and elastic fibers. The perimysium bound the bundles of muscle fibers together and frequently included blood vessels and nerves. As the superior head of the pterygoid muscle approached its insertion, sheaths of perimysium divided this head into smaller and smaller bundles of muscle fibers. In the inferior head, some of the perimysial sheaths and part of the epimysium were aponeurotic, and many muscle fibers attached to them. There were few such aponeurotic regions in the superior head. © 1996 Wiley-Liss, Inc.     

Influence of Passive Muscle Tension on Electromechanical Delay in Humans

Background

Electromechanical delay is the time lag between onsets of muscle activation and muscle force production and reflects both electro-chemical processes and mechanical processes. The aims of the present study were two-fold: to experimentally determine the slack length of each head of the biceps brachii using elastography and to determine the influence of the length of biceps brachii on electromechanical delay and its electro-chemical/mechanical processes using very high frame rate ultrasound.

Methods/Results

First, 12 participants performed two passive stretches to evaluate the change in passive tension for each head of the biceps brachii. Then, they underwent two electrically evoked contractions from 120 to 20° of elbow flexion (0°: full extension), with the echographic probe maintained over the muscle belly and the myotendinous junction of biceps brachii. The slack length was found to occur at 95.5 ± 6.3° and 95.3 ± 8.2° of the elbow joint angle for the long and short heads of the biceps brachii, respectively. The electromechanical delay was significantly longer at 120° (16.9 ± 3.1 ms; p<0.001), 110° (15.0 ± 3.1 ms; p<0.001) and 100° (12.7 ± 2.5 ms; p = 0.01) of elbow joint angle compared to 90° (11.1 ± 1.7 ms). However, the delay between the onset of electrical stimulation and the onset of both muscle fascicles (3.9 ± 0.2 ms) and myotendinous junction (3.7 ± 0.3 ms) motion was not significantly affected by the joint angle (p>0.95).

Conclusion

In contrast to previous observations on gastrocnemius medialis, the onset of muscle motion and the onset of myotendinous junction motion occurred simultaneously regardless of the length of the biceps brachii. That suggests that the between-muscles differences reported in the literature cannot be explained by different muscle passive tension but instead may be attributable to muscle architectural differences.