Characterization of skeletal muscles by MR imaging and relaxation times

Magnetic resonance (MR) images of three major flight muscles of chicks were obtained with surface coils using a 0.3 Tesla whole body imaging system (FONAR Beta 3000). The two fast muscles, pectoralis major (PM) and posterior latissimus dorsi (PLD), and a slow muscle, anterior latissimus dorsi (ALD), were identified in the axial, coronal, and sagittal images. The signal intensity (SI) of each muscle was electronically measured and its ratio to the background noise (S/N) was determined. Although visually the three muscles showed intermediate SI, the slow and fast muscles could be differentiated on the basis of their S/N values. These values were invariably higher in the slow muscles than in the fast muscles. To understand these differences, the muscles were excised and their mono- and multiexponential MR relaxation times (T1 and T2) were determined at 30 MHz. Multiexponential analysis enhanced the differences between the muscle types. With the sole exception of short T2, all relaxation components of the slow muscles were significantly longer than those of the fast muscles. These results suggest that elevation in the S/N, T1 and T2 values of muscles may not necessarily indicate a pathologic event, but may reflect the preponderance of slow fibers.     

Comprehensive analysis of titin protein isoform and alternative splicing in normal and mutant rats

Titin is a giant protein with multiple functions in cardiac and skeletal muscles. Rat cardiac titin undergoes developmental isoform transition from the neonatal 3.7 MDa N2BA isoform to primarily the adult 2.97 MDa N2B isoform. An autosomal dominant mutation dramatically altered this transformation. Titins from eight skeletal muscles: Tibialis Anterior (TA), Longissimus Dorsi (LD) and Gastrocnemius (GA), Extensor Digitorum Longus (ED), Soleus (SO), Psoas (PS), Extensor Oblique (EO), and Diaphram (DI) were characterized in wild type and in homozygous mutant (Hm) rats with a titin splicing defect. Results showed that the developmental reduction in titin size is eliminated in the mutant rat so that the titins in all investigated skeletal muscles remain large in the adult. The alternative splicing of titin mRNA was found repressed by this mutation, a result consistent with the large titin isoform in the mutant. The developmental pattern of titin mRNA alternative splicing differs between heart and skeletal muscles. The retention of intron 49 reveals a possible mechanism for the absence of the N2B unique region in the expressed titin protein of skeletal muscle.     

Coactivation of jaw muscles: recruitment order and level as a function of bite force direction and magnitude

The aim of this study was to obtain insight into the coactivation behaviour of the jaw muscles under various a priori defined static loading conditions of the mandible. As the masticatory system is mechanically redundant, an infinite number of recruitment patterns is theoretically possible to produce a certain bite force. Using a three-component force transducer and a feedback method, subjects could be instructed to produce a bite force of specific direction and magnitude under simultaneous registration of the EMG activity of anterior and posterior temporal, masseter and digastric muscles on each side. Forces were measured at the second premolars. Vertical, anterior, posterior, lateral and medial force directions were examined; in each direction force levels between 50 N and maximal voluntary force were produced. The results show that for all muscles the bite force-EMG relationship obeys a straight-line fit for forces exceeding 50 N. The relationship varies with bite force direction, except in the case of the digastric muscles. Variation is small for the anterior temporal and large for the posterior temporal and masseter muscles. The relative activation of muscles for a particular force in a particular direction in unique, despite the redundancy.     

Muscle growth and protein degradation during early development in chicks of fast and slow growing strains

1. Growth of breast and leg muscles and excretion of N tau methyl histidine in layer (slow growing) and broiler (fast growing) chicks were measured at five time intervals between 2 and 33 days of age. 2. The results indicate that muscles of the broiler chick grow faster than in layer chicks and that breast muscles of both strains grow faster than leg muscles in the first 2 weeks after hatching. 3. N tau methyl histidine excretion by layer chicks is higher than that by broilers relative to body weight, musculature and relative maturity at all ages examined. 4. The results suggest that faster growth of muscles is accompanied by a lower rate of protein degradation although at ages of less than 2 weeks differences in protein synthesis rates may also contribute to muscle growth.     

New titin (connectin) isoforms and their functional role in striated muscles of mammals: Facts and suppositions

This review summarizes results of our studies on titin isoform composition in vertebrate striated muscles under normal conditions, during hibernation, real and simulated microgravity, and under pathological conditions (stiff-person syndrome, post-apoplectic spasticity, dilated cardiomyopathy, cardiac hypertrophy). Experimental evidence for the existence in mammalian striated muscles of higher molecular weight isoforms of titin (NT-isoforms) in addition to the known N2A-, N2BA-, and N2B-titin isoforms was obtained. Comparative studies of changes in titin isoform composition and structure-functional properties of human and animal striated muscles during adaptive and pathological processes led to a conclusion about the key role of NT-isoforms of titin in maintenance of sarcomere structure and contractile function of these muscles.     

Differential effects of acute changes in cell Ca2+ concentration on myofibrillar and non-myofibrillar protein breakdown in the rat extensor digitorum longus muscle in vitro. Assessment by production of tyrosine and N tau-methylhistidine.

The influence of Ca2+ on myofibrillar proteolysis was evaluated in the isolated extensor digitorum longus muscle incubated in vitro with agents previously shown to increase the intracellular concentration of Ca2+. Myofibrillar proteolysis was evaluated by measuring the release of N tau-methylhistidine, and total proteolysis was evaluated by measuring tyrosine release by incubated muscles after the inhibition of protein synthesis with cycloheximide. Incubated muscles released measurable quantities of N tau-methylhistidine, and muscle contents of the amino acids remained stable over 2 h of incubation. The release of N tau-methylhistidine by incubated muscles was similar to its release by perfused rat muscle in response to brief starvation, indicating the integrity of the incubated muscles. Ca2+ ionophore A23187, dibucaine, procaine, caffeine and elevated K+ concentration increased lactate release by incubated muscles and decreased tissue contents of ATP and phosphocreatine to varying degrees, indicating the metabolic effectiveness of the agents tested. Only A23187 and dibucaine increased total cell Ca2+, and they increased tyrosine release. Caffeine and elevated [K+] increased neither cell Ca2+ nor tyrosine release; however, only A23187 and dibucaine increased tyrosine release significantly. On the other hand, these agents were without effect on myofibrillar proteolysis as assessed by N tau-methylhistidine release by incubated muscles and changes in tissue contents of the amino acid. In fact, some of the agents tested tended to decrease myofibrillar proteolysis slightly. These results indicate that acute elevation of intracellular Ca2+ is associated with increased breakdown of non-myofibrillar but not myofibrillar proteins. Because of this, the role of elevated Ca2+ in muscle atrophy in certain pathological states is questioned. The data also indicate that the breakdown of myofibrillar and non-myofibrillar proteins in muscle is regulated independently and by different pathways, a conclusion reached in previous studies with perfused rat muscle.     

Biomechanical model study of pelvic belt influence on muscle and ligament forces

Many patients with low back and/or pelvic girdle pain feel relief after application of a pelvic belt. External compression might unload painful ligaments and joints, but the exact mechanical effect on pelvic structures, especially in (active) upright position, is still unknown. In the present study, a static three-dimensional (3-D) pelvic model was used to simulate compression at the level of anterior superior iliac spine and the greater trochanter. The model optimised forces in 100 muscles, 8 ligaments and 8 joints in upright trunk, pelvis and upper legs using a criterion of minimising maximum muscle stress. Initially, abdominal muscles, sacrotuberal ligaments and vertical sacroiliac joints (SIJ) shear forces mainly balanced a trunk weight of 500N in upright position. Application of 50N medial compression force at the anterior superior iliac spine (equivalent to 25N belt tension force) deactivated some dorsal hip muscles and reduced the maximum muscle stress by 37%. Increasing the compression up to 100N reduced the vertical SIJ shear force by 10% and increased SIJ compression force with 52%. Shifting the medial compression force of 100N in steps of 10N to the greater trochanter did not change the muscle activation pattern but further increased SIJ compression force by 40% compared to coxal compression. Moreover, the passive ligament forces were distributed over the sacrotuberal, the sacrospinal and the posterior ligaments. The findings support the cause-related designing of new pelvic belts to unload painful pelvic ligaments or muscles in upright posture.     

Acute alterations in sodium flux in vitro lead to decreased myofibrillar protein breakdown in rat skeletal muscle.

Myofibrillar protein breakdown was evaluated by measuring the release of N tau-methylhistidine by isolated rat skeletal muscles or perfused rat muscles in the presence of a variety of agents known to affect Na+ flux. Total cell proteolysis was evaluated simultaneously by measuring tyrosine release by muscles after the inhibition of protein synthesis with cycloheximide. Treatment of muscles with the Na+ ionophore monensin or inhibitors of Na+-K+ ATPase (ouabain, digoxin or vanadate) decreased N tau-methylhistidine release by muscles by 21-35%. A phorbol ester (phorbol 12-myristate 13-acetate) as well as a synthetic diacylglycerol known to activate protein kinase C and a Na+/H+ antiport also decreased N tau-methylhistidine release by muscles. Removal of extracellular Na+ blocked the ability of these agents to attenuate N tau-methylhistidine release by muscles, suggesting that their effectiveness required a change in Na+ flux. In contrast with N tau-methylhistidine release by muscles, these agents, except for monensin, did not effect the release of tyrosine, suggesting that they attenuate specifically the breakdown of myofibrillar proteins. Overall these results indicate a link between Na+ and the regulation of protein breakdown in rat skeletal muscle, whereby an influx of Na+ can result in a decrease in myofibrillar proteolysis. Left unresolved is whether phospholipid hydrolysis is involved in this scheme.     

Power outputs of slow and fast skeletal muscles of mice

The purpose of this study was to contrast the frequency-power relationship of slow soleus and fast extensor digitorum longus (EDL) muscles to their frequency-force relationships and to investigate factors involved in the development of maximum power during a single contraction. Stimulation frequency-force and stimulation frequency-power relationships were determined for soleus and EDL muscles of the mouse for single contractions in situ at 35 degrees C. Power was measured during isovelocity shortening contractions with displacement through 10% of fiber length at the optimum velocity. Optimum velocity was defined as the shortening velocity for the generation of maximum power for a given stimulation frequency. Both force (N/cm2) and power (watts/kg) increased with stimulation frequency until a plateau was reached. For the frequency-force relationship, the curve for soleus muscles was merely shifted to the left of that for EDL muscles. In contrast, the power developed by EDL muscles was greater than that of soleus muscles (P less than 0.05) at each stimulation frequency. The higher power was a direct consequence of higher optimum velocities for EDL muscles compared with soleus muscles.     

Mutual influences of upper and lower extrimities during cyclic movements

The possibility of muscle activation of passive arm during its cyclic movements, imposed by active movements of contralateral arm or by experimenter was studied, as well as the influence of lower extremities cyclic movements onto arm muscles activity. In addition to that the activity of legs muscles was estimated in dependence on motor task condition for arms. Ten healthy supine subjects carried out opposite movements of arms with and without stepping-like movements of both legs. The experiment included three conditions for arm movements: 1) the active movements of both arms; 2) the active movements of one arm, when other entirely passive arm participated in the movement by force; 3) passive arm movement caused by experimenter. In the condition 2) additional load on active arm was applied (30 N and 60 N). In all three conditions the experiment was carried out with arms movements only or together with legs movements. The capability of passive moving arm muscles activation depended on increasing afferent inflow from muscles of contralateral arm was demonstrated. Emerging electrical activity was modulated in the arms movements cycle and depended on the degree of active arm loading. During combined active movements of arms and legs the reduction of activity in the flexor muscles of shoulder and forearm was observed. Concomitant arms movements increased the magnitude ofelectromiographic bursts during passive stepping-like movements in the most of recorded muscles, and the same increasing was only observed in biceps femoris and tibialis anterior muscles during active legs movement. The increasing of loading of one arm caused essential augmentation of EMG-activity in the majority of recording legs muscles. The data obtained are the additional proof of existence of functionally significant neuronal interaction both between arms and between upper and lower extremities, which is evidently depend on the intraspinal neuronal connections.     

The minimum number of muscles to control a chain of joints with and without tenodeses, arthrodeses, or braces--application to the human finger

While the underlying principles of controlling a single joint have been well described, the principles of simultaneously controlling multiple joints have not been comprehensively addressed in the literature of reconstructive hand surgery. This article analyzes (1) how many muscles are minimally required to fully control a chain of joints with in total N Degrees of Freedom (DoF), and (2) to what degree tenodeses, arthrodeses or braces can reduce the required number of muscles. It is demonstrated by mathematical analysis and illustrated by examples that the minimal number of muscles to control a chain of N DoF is N + 1. The number of muscles required for control can be reduced by mechanisms that reduce the number of DoF in the chain. (i) An arthrodesis is a permanent surgical fixation of a joint. An arthrodesis eliminates as many DoF in the chain as the arthrodized joints contributed. (ii) Tenodeses are coordinative tendon constructions. Each independent tenodesis eliminates one DoF from the chain. (iii) Braces are removable external supports. They eliminate as many DoF for muscle control as they immobilize. These principles are applied to illustrate the fundamental importance of tendinous structures in control in the human finger. Being able to determine the minimum number of muscles needed for multiarticular control gives additional knowledge in the design of functional reconstruction.     

Mechanically simulated muscle forces strongly stabilize intact and injured upper cervical spine specimens

Although muscles are assumed to be capable of stabilizing the spinal column in vivo, they have only rarely been simulated in vitro. Their effect might be of particular importance in unstable segments. The present study therefore tests the hypothesis that mechanically simulated muscle forces stabilize intact and injured cervical spine specimens. In the first step, six human occipito-cervical spine specimens were loaded intact in a spine tester with pure moments in lateral bending (+/- 1.5 N m), flexion-extension (+/- 1.5 N m) and axial rotation (+/- 0.5 N m). In the second step, identical flexibility tests were carried out during constant traction of three mechanically simulated muscle pairs: splenius capitits (5 N), semispinalis capitis (5 N) and longus colli (15 N). Both steps were repeated after unilateral and bilateral transection of the alar ligaments. The muscle forces strongly stabilized C0-C2 in all loading and injury states. This was most obvious in axial rotation, where a reduction of range of motion (ROM) and neutral zone to <50% (without muscles=100%) was observed. With increasing injury the normalized ROM (intact condition=100%) increased with and without muscles approximately to the same extend. With bilateral injury this increase was 125-132% in lateral bending, 112%-119% in flexion-extension and 103-116% in axial rotation. Mechanically simulated cervical spine muscles strongly stabilized intact and injured cervical spine specimens. Nevertheless, it could be shown that in vitro flexibility tests without muscle force simulation do not necessarily lead to an overestimation of spinal instability if the results are normalized to the intact state.     

Accelerated contractile function and improved fatigue resistance of calf muscles in newborn piglets with IUGR

Asymmetrical intrauterine growth restriction is denoted by disproportional reduction of muscle mass compared with body weight reduction. However, effects on contractile function or tissue development of skeletal muscles were not studied until now. Therefore, isometric force output of serial-stimulated hindlimb plantar flexors was measured in thiopental-anesthetized normal weight (NW) and intrauterine growth-restricted (IUGR) 1-day-old piglets under conditions of normal, reduced (aortic cross clamping), and reestablished (clamp release) blood supply (measured by colored microspheres technique). Furthermore, muscle fiber type distribution was determined after histochemical staining, specific muscle force of the plantar flexors [quotient from absolute force divided by muscle mass (N/g)] was calculated, and glycogen content and morphometric data of the investigated muscles were estimated. Regional blood flow of hindlimb muscles was similar in NW (6 +/- 2 ml. min(-1). 100 g(-1)) and IUGR piglets (8 +/- 1 ml. min(-1). 100 g(-1)). Isometric muscle contractions induced a marked increase in regional blood flow of 4.1-fold in NW and 5-fold in stimulated hindlimb muscles of IUGR piglets (baseline blood flow). Specific force of NW piglet muscles (5.2 +/- 0.2 N/g) was significantly lower than IUGR piglet muscles (6.1 +/- 0.6 N/g; P < 0.05). Isometric muscle contractions (NW: 32.7 +/- 4.7 N; IUGR: 21.7 +/- 4.0 N) resulted in a higher rate of force decrease in the calf muscles of NW animals compared with IUGR piglets (8 +/- 2 vs. 3 +/- 1%; P < 0. 01). Functional restoration of contractile performance after hindlimb recirculation was nearly complete in IUGR piglets (98 +/- 1%), whereas in NW piglets a deficit of 9 +/- 3% was found (P < 0. 01). Muscle fiber type estimation revealed an increased proportion of type I fibers in flexor digitalis superficialis and gastrocnemius medialis in IUGR piglets (P < 0.05). These data clearly indicate that contractile function is accelerated in newborn IUGR piglets.     

Pulmonary ventilation, blood gases, and blood pH after training of the arms or the legs.

In two groups of young healthy subjects who performed arm training (N = 5) and leg training (N = 5), respectively, the respiratory adaptation to submaximal exercise with trained and nontrained muscle groups was compared by measurement of the ventilatory equivalent (Ve/Vo2, pH, and blood gases (Pco2, Po2, and So2) in arterial blood and in venous blood from exercising extremities. After training Ve/Vo2 was significantly reduced during exercise with trained muscles, but unchanged during exercise with nontrained muscles. The reduction in Ve/Vo2 was closely related to a less pronounced increase in heart rate and in arterial lactate content, but showed no quantitative correlation to changes in arterial adaptations in trained muscles are mainly responsible for the reduction in Ve/Vo2. After training during exercise with trained as well as nontrained muscles a shift to the right of the blood oxygen dissociation curve occurred as extremities was lower while corresponding Po2 was higher.     

Inhibitory effect of genital photoreceptors on the activity of abdominal motoneurones in Papilio xuthus (Lepidoptera,Papilionidae)

The inhibitory effect of butterfly genital photoreceptors on the activities of abdominal motoneurones is described. In two (N1, N3) of the six lateral nerves (N1–6) belonging to the last abdominal ganglion, spontaneous motoneurone activity was inhibited by illumination of the genital photoreceptors. N1 and N3 innervate the ventral longitudinal muscles. N2 and N4, which supply branches to the spiracular muscles, were not inhibited. The results allow some of the properties of the circuits in the abdominal nervous system supplied by the genital photoreceptors to be inferred. Some possible functions of the photoreceptors are discussed.     

Seasonal changes in the composition of titin isoforms in muscles of hibernating ground squirrels

An electophoretic study of changes in the content of intact titin isoforms, N2B-, N2BA-, N2A-titins and T2 in skeletal and cardiac muscles of ground squirrel (Spermophillus undulatus) is made in different periods: summer activity, autumnal activity, hibernation, arousal, and winter activity. In atria and ventricles of ground squirrels in the period of autumnal activity an increase (by ～1.5 times) in the N2BA to N2B ratio was observed, in comparison with that in cardiac muscle in summer activity. During hibernation, the decrease in the relative content of N2B-, N2BA-titins and T2 in cardiac muscle as well as of N2A-titin and T2 in skeletal muscles was determined against the background of preservation of the relative amount of intact titin isoforms. At waking of ground squirrels and in a short period of winter activity, a rapid restoration of the content of N2B-, N2BA-, N2A-titisns and T2 in muscles was observed. In the myocardium of hibernating, waking ground squirrels and of those during winter activity the increased N2BA to N2B ratio was retained. The changes in the titin content are discussed in the aspect of adaptation of ground squirrels to hibernation.     

Structure and direction of jaw adductor muscles as herbivorous adaptations in <Emphasis Type="Italic">Neotoma mexicana</Emphasis> (Muridae,Rodentia)

The herbivorous adaptations of the jaw adductor muscles in Neotoma mexicana were clarified by a comparative study with an unspecialized relative, Peromyscus maniculatus. In P. maniculatus, the anterior part of the deep masseter arises entirely from the lateral side of an aponeurosis, i.e., superior zygomatic plate aponeurosis, whereas N. mexicana has an additional aponeurosis for this part of the muscle, and the fibers attach on both sides of the superior zygomatic plate aponeurosis. Although the structure of the temporalis muscle is nearly identical in the two genera, a clear aponeurosis of origin occurs only in N. mexicana. These characteristics allow fibrous tissues to be processed with a large occlusal force. The deep masseter, internal pterygoid, and external pterygoid muscles of N. mexicana incline more anterodorsally than those of P. maniculatus. The transverse force component of these muscles relative to whole muscle force is smaller in N. mexicana than in P. maniculatus, with the exception of the internal pterygoid. The anterior part of the temporalis muscle of N. mexicana is specialized to produce occlusal pressure. These findings suggest that in N. mexicana a large anterior force is required to move the heavy mandible, due to the hypsodont molars, against frictional force from food, and that the posterior pull of the temporalis, which adjusts the forward force by the other jaw adductor muscles to a suitable level, need not be large for the mandibular movement.     

Passive and active mechanical properties of the superficial and deep digital flexor muscles in the forelimbs of anesthetized Thoroughbred horses

The superficial (SDF) and deep digital flexor (DDF) muscles are critical for equine forelimb locomotion. Knowledge of their mechanical properties will enhance our understanding of limb biomechanics. Muscle contractile properties derived from architectural-based algorithms may overestimate real forces and underestimate shortening capacity because of simplistic assumptions regarding muscle architecture. Therefore, passive and active (=total - passive) force-length properties of the SDF and DDF muscles were measured directly in vivo. Muscles from the right forelimbs of four Thoroughbred horses were evaluated during general anesthesia. Limbs were fixed to an external frame with the muscle attached to a linear actuator and load cell. Each muscle was stretched from an unloaded state to a range of prefixed lengths, then stimulated while held at that length. The total force did not exceed 4000 N, the limit for the clamping device. The SDF and DDF muscles produced 716+/-192 and 1577+/-203 N maximum active isometric force (F(max)), had ascending force-length ranges (R(asc)) of 5.1+/-0.2 and 9.1+/-0.4 cm, and had passive stiffnesses of 1186+/-104 and 1132+/-51 N/cm, respectively. The values measured for F(max) were much smaller than predicted based on conservative estimates of muscle specific tension and muscle physiological cross-sectional area. R(asc) were much larger than predicted based on muscle fiber length estimates. These data suggest that accurate prediction of the active mechanical behavior of architecturally complex muscles such as the equine DDF and SDF requires more sophisticated algorithms.     

Contributions of slow and fast muscles of triceps surae to a cyclic movement

The relative contribution of synergistic muscles has been studied during pedalling on a bicycle. The electromyographic (EMG) activity of the different components of triceps surae (namely soleus or SOL and medial gastrocnemius or MG) has been recorded and analyzed for increasing pedalling speed performed against increasing resistance. The results indicate that SOL IEMG (integrated EMG) increases linearly (y = 2x-12.1; r = 0.98) with increasing load (10-70 N) at constant speed (60 rpm), whereas no change is noted in MG IEMG below 40 N. In contrast, when the pedalling speed is increased (from 30 to 170 rpm) at constant load, MG IEMG shows the largest increase. Furthermore, although in both muscles EMG activity appears earlier in the movement with increases in load and/or speed, the delay between the onset of both EMGs remains unchanged at constant speed and synchronization of MG with SOL is only observed when speed is increased above 140 rpm. These results suggest that the different muscles of the triceps surae make specific contributions to the development of the mechanical tension required to maintain or increase the speed of movement.