Macroscopic-microscopic characterization of the passive mechanical properties in rat soleus muscle.

The purpose of the study was to investigate changes in passive mechanical properties of the soleus muscle of the rat during the first year of life. These mechanical changes were quantified at a macroscopic (whole muscle) and a microscopic level (fiber) and were correlated with biochemical and morphological properties. Three passive mechanical tests (a relaxation test, a ramp stretch test and a stretch release cycle test) with different amplitudes and velocities were performed on isolated soleus muscles and fibers in rats at ages 1 (R1), 4 (R4) and 12 (R12) months. Mechanical parameters (dynamic and static forces, stresses and normalized stiffness) were recorded and measured. The morphological properties (size of fibers and muscles) for the three groups of rats were assessed by light microscopy which allowed us to observe the evolution of the fiber type (I, IIc and IIa) in the belly region and along the longitudinal axis of the muscle. In addition, biochemical analyses were performed at the level of the whole muscle in order to determine the collagen content. The results of the passive mechanical properties between the macroscopic (muscle) and microscopic (fiber) levels showed a similar evolution. Thus, an increase of the dynamic and static forces appeared between 1 and 4 months while a decrease of the passive tension occurred between 4 and 12 months. These mechanical changes were correlated to the morphological properties. In addition, the size of the three fibers type which grew with age could explain the increase of forces between 1 and 4 months. Furthermore, the biochemical analysis showed an increase of the collagen content during the same period which could also be associated with the increase of the passive forces. After 4 months, the passive tension decreased while the size of the fiber continued to increase. The biochemical analysis showed a decrease of the collagen content after 4 months, which could explain the loss of passive tension in the whole muscle. Concerning the similar loss at the fiber level, other assumptions are required such as a myofibril loss process and an increase of intermyofibrillar spaces. The originality of this present study was to compare the passive mechanical properties between two different levels of anatomical organization within the soleus muscle of the rat and to explain these mechanical changes in terms of biochemical and morphological properties.

Relationship between muscle myosin isoforms and contractile features in rabbit fast-twitch denervated muscle.

The effects of 8-day-old rabbit fast-twitch gastrocnemius denervation on the type of myosin isoforms and on contractile features (maximum velocity Vmax and contraction time (CT) of the muscle were followed between 15 and 60 days postnatal. The myosin isoforms and the Vmax and CT values of the denervated gastrocnemius displayed large changes during this period. These changes, which led at 2 months postnatal to a muscle displaying the properties of a slow-twitch muscle did not occur in synchrony: complete conversion to slow-type myosin isoforms occurred only at 60 days postnatal, whereas complete conversion to slow-twitch Vmax and CT values occurred as soon as 35 days postnatal. The results address a new question concerning the relationship between muscle myosin and contractile features.

Effects of long-term spaceflight on mechanical properties of muscles in humans.

The effects of long-term spaceflight (90-180 days) on the contractile and elastic characteristics of the human plantarflexor muscles were studied in 14 cosmonauts before and 2-3 days after landing. Despite countermeasures practiced aboard, spaceflight was found to induce a decrease in maximal isometric torque (17%), whereas an index of maximal shortening velocity was found to increase (31%). In addition, maximal muscle activation evaluated during isokinetic tests decreased by 39%. Changes in musculotendinous stiffness and whole joint stiffness were characterized by means of quick-release movements and sinusoidal perturbations. Musculotendinous stiffness was found to be increased by 25%. Whole joint stiffness decreased under passive conditions (21%), whereas whole joint stiffness under active conditions remained unchanged after spaceflight (-1%). This invariance suggests an adaptive mechanism to counterbalance the decrease in stiffness of passive structures by an increased active stiffness. Changes in neural drive could participate in this equilibrium.

Surface electromyogram power spectrum in human quadriceps muscle during incremental exercise.

Spectral electromyographic (EMG) changes in human quadriceps muscles were studied to reinvestigate discrepant results concerning mean power frequency (MPF) changes during dynamic exercise. An incremental test consisting of a quasi-linear increase in mechanical power on a bicycle ergometer (for 20-100% of maximal aerobic power) was performed by forty subjects. During this test, surface EMGs from the quadriceps muscles showed that EMG total power (PEMG) increased with a curvilinear pattern for every subject, whereas MPF kinetics varied from one subject to another. PEMG changes had the same shape, which would lead to disappointing results in terms of discrimination between subjects. The ability of normalized MPF kinetics to define significant clusters of subjects was tested using a principal component analysis. This analysis led to the projection of all experiments onto a plane and revealed a relevant grouping of MPF profiles. Differences in MPF kinetics between clusters are interpreted in terms of various possibilities of balance between physiological events leading to an increase or a decrease in MPF.

Surface EMG power spectrum and intramuscular pH in human vastus lateralis muscle during dynamic exercise.

The relationship between intramuscular pH and the frequency components of the surface electromyographic (EMG) power spectrum from the vastus lateralis muscle was studied in eight healthy male subjects during brief dynamic exercise. The studies were carried out in placebo control and metabolic alkalosis induced by oral administration of NaHCO3. At the onset of exercise, blood pH was 0.08 units higher in alkalosis compared with placebo. Muscle lactate accumulation during exercise was higher in alkalosis (32 +/- 5 mmol/kg wet wt) than in placebo (17 +/- 4 mmol/kg wet wt), but no difference in intramuscular pH was found between the two conditions. The EMG power spectrum was shifted toward lower frequencies during fatigue in the control condition (10.1 +/- 0.9%), and these spectral shifts, evaluated from changes in the mean power frequency (MPF) of the EMG power spectrum, were further accentuated in alkalosis (19 +/- 2%). Although the changes in frequency components of EMG correlated with muscle lactate accumulation (r = 0.68, P less than 0.01), no direct relationship with muscle pH was observed. We conclude that alkalosis results in a greater reduction in MPF associated with a higher muscle lactate accumulation. However, the good correlation observed between the two variables is not likely causative, and a dissociation between intramuscular pH and the increase in the low-frequency content of EMG power spectrum appears during muscle fatigue.

Surface electromyogram during voluntary contraction: processing tools and relation to physiological events.

Surface electromyography (SEMG) has been used extensively in the last years in a variety of applications, including muscle function assessment, pathology identification, ergonomics, pattern analysis, or population characterization. Advanced processing methods, especially in the spectral domain, provide the research worker with more and more precise and user-friendly tools for signal characterization, analysis, and classification. The use of such sophisticated tools requires many assumptions on signal characteristics, and the wide variety of computing options related to each processing method makes it difficult to compare the results of different works when these options are omitted in the reports or improperly applied. This work first aims at taking stock of the various processing methods which have emerged in the last years around surface electromyography: signal acquisition, random feature extraction, time and spectral parameter determination, statistical tests application. The main methods are briefly explained and discussed, then variations between apparently equivalent methods are pointed out, necessary hypotheses are underlined, and the use of such methods in SEMG processing is shown with respect to the more recent works. A second section shows how authors deal with parameters extracted from SEMG in order to relate them to physiological modifications (force, fiber type, fiber environment).

Endotoxins modify muscle fatigue characteristics.

To test the hypothesis that endotoxins can directly modify muscle fatigue characteristics, in vitro experiments were performed on rat muscles 48 hours after injection of lipopolysaccharides (LPS) from Klebsiella pneumoniae. Resistance to fatigue was quantified by measuring tension production during repetitive electrical stimulation of the isolated epitrochlearis muscle. LPS treatment did not significantly modify initial force production whereas fatigability of the muscle was increased. This in vitro preparation should be used for testing antifatigue drugs.

In vivo force-velocity relation of human muscle: a modelling from sinusoidal oscillation behaviour.

Isokinetic tests performed on human muscle in vivo during plantar flexion contractions lead to torque-angular velocity relationships usually fitted by Hill's equation expressed in angular terms. However, such tests can lead to discrepant results since they require maximal voluntary contractions performed in dynamic conditions. In the present study, another way to approach mechanical behaviour of a musculo-articular structure was used, i.e. sinusoidal oscillations during sub-maximal contractions. This led to the expression of (i). Bode diagrams allowing the determination of a damping coefficient (B(bode)); and (ii). a viscous parameter (B(sin)) using an adaptation of Hill's equation to sinusoidal oscillations. Then torque-angular velocity relationships were predicted from a model based on the interrelation between B(bode) and B(sin) and on the determination of optimal conditions of contraction. This offers the possibility of characterizing muscle dynamic properties by avoiding the use of isokinetic maximal contractions.

Models of skeletal muscle to explain the increase in passive stiffness in desmin knockout muscle.

Absence of desmin in skeletal muscle was found to induce an increase in passive stiffness. The present study aimed at developing rheological models of passive muscle to explain this stiffening. Models were elaborated by using experimental data depicting muscle viscoelastic behaviour. The experimental protocol included stepwise extension tests applied on control and desmin knockout soleus muscles from mice. Linear and non-linear models were composed of elastic and viscous elements. They were constructed with the aim at taking the presence or absence of desmin into account by simulating desmin as an elastic element. Furthermore, associated adaptation of connective tissues in absence of desmin was modelled as an additional elastic element. Differences in passive behaviour induced by absence of desmin were predicted by using a linear model and a non-linear one. The non-linear model was selected because: (1) it is able to predict experimental viscoelastic kinetics accounting for the increase in passive stiffness in muscles lacking desmin, (2) its design is consistent with morphological data, and (3) stiffness characteristics of its elements are in accordance with the literature. Finally, this modelling approach demonstrates that both absence of desmin and adaptation of connective tissue are required to explain the increase in passive stiffness in desmin knockout muscles.

Viscous properties of human muscle during contraction.

The purpose of this study was to determine viscous properties of human muscle during plantarflexion efforts. Experiments were performed on 17 subjects with an ankle ergometer allowing sinusoidal oscillations during isometric contractions and isokinetic movements. Sinusoidal oscillations led to the expression of (i) Bode diagrams of the musculo-articular system allowing the determination of a damping coefficient (Bbode); and (ii) a viscous coefficient (Bsin) using an adaptation of Hill's equation to sinusoidal oscillations. Isokinetic movements led to torque-velocity relationships. They showed a fall in torque associated to an increase in angular velocity what was quantified by calculating a damping coefficient (Biso). Both experiments gave consistent results indicating that Bbode was the lowest viscous parameter. This difference is discussed in terms of (i) "analog" viscosity originating from muscle cross-bridges; and (ii) real mechanical damping of passive structures.

Series elasticity in frog sartorius muscle subjected to stretch-shortening cycles.

Changes in stiffness during stretch-shortening cycles were studied by applying prestretches of different rates and amplitudes on the tetanized sartorius muscle of the frog and measuring series elastic component characteristics during a subsequent quick release. Series elastic component stiffness was found to be dependent on the mechanical parameters of the stretching phase, and the so-called 'Cavagna effect' (increase in compliance) was rarely observed. The results are discussed in terms of improvement in muscle efficiency. Thus, an increase in stiffness during force generation will favour a better transmission of force and will reduce the coupling time whereas the opposite evolution during SEC recoil will allow a better release of potential energy.

A mechanical device for studying mechanical properties of human muscles in vivo.

A mechanical device, primarily devoted to biomechanical studies of human calf muscles during microgravity experiments is presented. It allows investigation of both contractile and visco-elastic properties of musculo-articular systems using, respectively, isokinetic movements, quick-release tests and sinusoidal perturbations. This device is a specifically designed ergometer associated to an experimental protocol designed for pre- and post-flight tests. The protocol was evaluated on 22 healthy subjects and typical results are briefly presented. Preliminary results are discussed in terms of agreement with currently available data and a detailed evaluation of test-retest measurements is provided for quick-release experiments. Complementary investigations are suggested and potential fields of research are indicated.

Changes in elastic characteristics of human muscle induced by eccentric exercise.

The effect of an eccentric strength training programme on the muscular series elastic component (SEC) was studied on the flexors of the human elbow. The characteristics of the SEC were determined using an in situ technique derived from methods commonly used on isolated muscles. The results were expressed in terms of compliance-force and tension-extension relationships. These relationships indicate a sharp increase in compliance when tension decreases. Furthermore, for a given value of tension, the SEC compliance of the trained muscles is found to be lower than that of the untrained muscles. These results are discussed in relation to the active and passive parts of the SEC.

Reflex assessment of a reciprocal inhibition between bifunctional ago-antagonist muscles.

Neuronal connections between the peroneus longus and the soleus, ago-antagonist bifunctional muscles, were investigated by reflex analysis. The peroneus longus H-reflex, obtained by stimulating the common peroneal nerve, was conditioned by a posterior tibial nerve stimulation. This conditioning stimulus induced an early inhibition of the peroneus longus H-reflex and the degree of inhibition increased with the conditioning stimulus intensity. The soleus H-reflex, solicited by stimulating the posterior tibial nerve, was conditioned by passive adduction movements. These movements were passive quick-release movements achieved by a specific device containing an electromagnet and stiff springs. The passive adduction movement induced both a myotatic reflex in the stretched muscles and an early inhibition of the test soleus H-reflex. This inhibition is most likely due to the activation of the primary spindle afferents of the stretched muscles. Thus we propose an antagonist neuronal scheme between peroneus longus and soleus muscles in spite of their agonism in plantarflexion.

Surface myoelectric signals during ergocycle exrcises at various mechanical powers and pedalling rates.

The surface myoelectric (ME) signal from the vastus lateralis muscle was studied during ergocycling at various mechanical powers (MPs) and pedalling rates (PRs). The envelope of the signal was described using a power spectral density function estimate at very low frequency. A highly smoothed burst pattern was found independently at 80 rpm on MP. The envelope presence in surface ME signals was demonstrated to affect the total band power spectrum estimate slightly. In addition, the total band power spectrum changes with MPs and PRs were described using the root mean square value and the mean power frequency. Total band power spectrum was enhanced and shifted toward the low frequencies as PR increased. It was shifted toward the high frequencies as MP increased. These changes were attributed to the progressive recruitment of fast fibers.

Duchenne muscular dystrophy quantification: a multivariate analysis of surface EMG.

The paper describes a method of quantifying Duchenne muscular dystrophy which is examiner independent and uses surface electromyographic signals (EMG). A standardised protocol is proposed. Spectral parameters are first computed from digitised EMG, then a polynomial model is deduced from the evolution of each parameter. A discriminant analysis between healthy and DMD subjects leads to the determination of a discriminant plane and a level of sickness index.

Musculo-tendinous and joint elastic characteristics during elbow flexion in children.

OBJECTIVE: The aim of the present research was to determine joint stiffness and musculo-tendinous stiffness in children. BACKGROUND: No reliable data was available on differences in such parameters when comparing children and adults. METHODS: Sinusoidal perturbations and quick-released movements were applied to the right elbow during flexion to determine joint stiffness and musculo-tendinous stiffness, respectively. For normalization purposes, stiffness indexes were calculated as the slope of the relationship between stiffness and torque. RESULTS: Main results were that musculo-tendinous stiffness indexes obtained in children were similar to adult data deduced from the literature. On the other hand, joint stiffness indexes obtained in children were in the range 0.9-3.8 rad(-1) and were lower than those reported in the literature for adults (between 4 and 8.5 rad(-1)). CONCLUSION: Since normalized musculo-tendinous stiffness is similar in children and in adults, it can be hypothesized that, during growth, increase in ability to produce force parallels elastic tissue development. RELEVANCE: Stiffness parameters could be used, in the context of other functional tests, to characterize changes in muscle properties in neuromuscular disorders when elastic tissues are supposed to be altered.

Muscle stretch used as conditioning stimulus for assessing a reciprocal inhibition.

The disynaptic Ia-alpha inhibition between the tibialis anterior and the soleus muscles is now well known. This neuronal organization has been established thanks to the analysis of the Hoffmann reflex (H reflex) changes following an electrical stimulation of the antagonist muscle nerve. In some cases, anatomical constraints impede the use of this classical technique for assessing a reciprocal Ia inhibition between muscles. Furthermore, an electrical stimulus solicits the primary spindle afferents in conditions which are very different from their natural stimulus: the muscle stretch. Thus we have undertaken to analyse the changes of a soleus H reflex following a rapid stretch of the ankle dorsiflexors and therefore of their prime-mover: the tibialis anterior. The mechanical perturbations were imposed with a strong initial acceleration and a limited angular displacement thanks to an original device including electromagnet and spring. This technique, derivated from the quick release technique, was applied on relaxed muscles to avoid a co-contraction phenomenon. 45 to 50 ms after the initiation of the mechanical perturbation an early reflex response was observed on the stretched tibialis anterior. The area of this first reflex response was related to the initial acceleration of the passive dorsiflexion. Several arguments are presented in favour of the myotatic origin of this reflex component. The passive dorsiflexion used as a conditioning stimulus led to a strong and longlasting inhibition of the test soleus H reflex. The early inhibition of the soleus H reflex was observed only about 18 ms after the conditioning stimulus.(ABSTRACT TRUNCATED AT 250 WORDS)

The ankle ergometer: a new tool for quantifying changes in mechanical properties of human muscle as a result of spaceflight.

A mechanical device for studying changes in mechanical properties of human muscle as a result of spaceflight is presented. Its main capacities are to allow during a given experiment investigation of both contractile and visco-elastic properties of a musculo-articular complex using respectively isometric contractions, isokinetic movements, quick-release tests and sinusoidal perturbations. This device is a motor driven ergometer associated to an experimental protocol designed for pre- and post-flight experiments. As microgravity preferentially affects postural muscles, the apparatus was designed to test muscle groups crossing the ankle joint. Three subjects were tested during the Euromir' 94 mission. Preliminary results obtained on the european astronaut are briefly reported. During the next two years the experiments will be performed during six missions.