Chantal Pérot

Changes in stiffness induced by hindlimb suspension in rat Achilles tendon

Abstract The aim of this study was to measure the effects of hindlimb suspension on mechanical properties of the rat Achilles tendon. Adult male Wistar rats were randomly assigned to groups to be either suspended, or a control. After 21 days, Achilles tendons were removed for mechanical analysis. Classical tests of tensile performance were made, and mechanical parameters were derived from a stress-strain relationship. The tendons of animals that had been suspended presented values for maximal stress and tangent modulus which were 37.5% (P < 0.01) and 41% (P < 0.01), respectively, lower than the tendons of the control rats. In a similar way, the energy absorption capacity had largely decreased in animals that had been suspended. However, the maximal strain was similar in the two groups. These results showed that hindlimb suspension in rats has an important detrimental effect on mechanical properties of the Achilles tendon. Differences in tendon stiffness obtained here, along with those found by other investigators, encourage the hypothesis that homeostatic responses of soft tissues are due to changes in limb loadings. This study may be useful in providing a better understanding of the adaptation of human skeletal muscle when exposed to microgravity.

Differential effect of knee extension isometric training on the different muscles of the quadriceps femoris in humans

Abstract This study determined the effects of a short period of knee isometric training on the quadriceps muscles accessible to surface electromyography (EMG). For this purpose, a training (n=9) and a control (n=7) group were tested on five identical occasions at 1 week intervals during 4 weeks. The training group exercised three times a week by making isometric knee extensions at 80% of the maximal voluntary contraction (MVC). During the test sessions, maximal and submaximal torque and associated activations of the rectus femoris (RF), vastus lateralis (VL) and vastus medialis (VM) muscles were analysed. As a result of training, differences between MVC values of the two groups were highly significant (P < 0.001), whereas only RF-EMG showed significant differences (P < 0.05). The VL and VM did not present any significant changes in maximal activation. The EMG–torque relationships were analysed individually before and after the training period. For the control subjects, EMG–torque relationships did not present significant changes while for the training group, these relationships showed a significant increase in RF, VL, and VM maximal activation in 6, 6 and 4 subjects, respectively, and a significant decrease in 1, 2 and 5 subjects, respectively. In almost all cases, a significant downward shift of the relationship was observed. This study confirmed that the parts of the quadriceps muscle tested present different adaptation capacities and demonstrate inter-individual variability in the strategies used to enhance muscle strength. In conclusion, to analyse the neural effects resulting from training in a large and compartmentalized muscle like the quadriceps femoris, it is desirable to take into account each muscle independently. Moreover, we suggest that overall results obtained from the experiment population should be completed by an analysis on individuals.

Stiffness changes and fibre type transitions in rat soleus muscle produced by jumping training

Rat soleus muscles were overloaded with intent to induce a relative increase in fast fibres and modifications in muscular stiffness. The overloading technique was a training period consisting of an 11-week vertical jump programme. The method of controlled releases was used to obtain tension/extension curves characterizing the elastic behaviour of the soleus. Fibre typing was made by myofibrillar adenosine 5′-triphosphatase staining. With regard to a control group, training resulted in a relative decrease in type I fibres for the benefit of type II fibres. Training also induced a decrease in muscle stiffness as attested notably by significant differences in maximal extension. These results are interpreted in terms of modifications occurring in the active fraction of the so-called series elastic component.

Quantification of T- and H-responses before and after a period of endurance training

SummaryTendon (T-) and Hoffmann (H-) responses in the soleus muscle were quantified either separately or in association to compare the mononeurons activated and to study their changes after a period of endurance training. In a first experiment T- and H-responses of the same amplitude were compared: the electrical stimulus (inducing the H-response) and the Achilles tendon tap (inducing the T-response) were associated so that the T-response firstly was concomitant with the H-response, and secondly shifted 10 ms forward or back compared to the H-response. From the study of these combined reflexes we would suggest that the same motoneurons are involved in T- or H-responses of the same amplitude. In a second experiment the maximal H-responses, the T-responses and maximal aerobic power (Waer,max) were measured on 20 subjects before and after a period of endurance training. For 75% of the subjects the Waer,max and the reflex parameters (T or H) varied in the same direction: most of them exhibited higher values of both Waer,max and reflex amplitudes while the others had Waer,max and reflex values hardly modified or decreased. The different effects of the training period could reflect the heterogeneity of the subjects status and involvement in sport. In most cases the T: Hmax ratios were also influenced, reflecting the fact that T- and H-responses were not identically affected by training. Thus it is suggested that an endurance training programme can influence not only the excitability of the motoneurons but also the response of the muscle receptors to stretch. An interpretation in terms of a change of spindle receptivity and/or a change in their recruitment due to a greater stiffness of the trained muscles is suggested.

Age-Related Changes in Twitch Properties of Plantar Flexor Muscles in Prepubertal Children

The twitch of the triceps surae muscle (TS), which characterizes the contractile properties independently of volition, differs in amplitude, but not in time course, when evoked in pre or postpubertal children. The aim of the present study was to compare the TS twitch contractile properties in prepubertal children (7 to 11 y). M-wave and twitch were recorded at rest by supramaximal electrical stimulations of the posterior tibial nerve. Twitches were characterized by peak torque (Pt), contraction time (CT), half relaxation time (HRT), and rate of torque development (dPt/dt). Electromechanical delay (EMD) was quantified with regard to the TS M-wave onset. Pt values increased significantly with the age of the prepubertal children but remained lower than that for adult subjects. CT and HRT values did not change with age. Thus, dPt/dt increased significantly between the 7-year-old and the 11-year-old children but remained significantly lower than that for adults. Despite EMD values decreased with age, they remained significantly higher than those of adult subjects. These results confirmed the link between growth processes and the increase in twitch torque for prepubertal children within a limited range of age. However, the time-course characteristics were not affected by age. The increase in dPt/dt and the decrease in the EMD could be used as indirect indicators of changes in contractile kinetics and in musculo-tendinous stiffness with the age of the prepubertal children. The age-related relationships established by this study will serve as reference values for clinical testing of the TS performances in relation to muscle disease or disuse.

Effects of stretch-shortening cycle training on mechanical properties and fibre type transition in the rat soleus muscle

The effects of exercise training on mechanical properties and fibre type transitions have been investigated in rat soleus muscle. The exercise was a repetition of stretch-shortening cycles. A method of dual controlled releases was applied to obtain tension/extension curves, which characterize the elastic behaviour of the series elastic component (SEC), and the force/velocity relationship characterizing the contractile elements. Other contractile measurements included: contraction time (tc), half-relaxation time (t1/2) and twitch/tetanus ratio (Pt/Po). Additionally, the muscle fibre type composition was determined by a classical histochemical method. A 12-week period of training induced a significantly higher percentage of fast-twitch fibres and a lower percentage of slow-twitch fibres (P<0.01). This fibre adaptation led to a significant (P<0.01) decrease in tc and an increase in maximum shortening velocity (Vmax). An increase in compliance of the SEC was also observed. This elastic adaptation is interpreted in terms of modification of the active components of the SEC. All the histochemical and mechanical data presented in this study show that rat soleus muscles trained by stretch-shortening cycles acquired faster characteristics. Thus the results confirm that a slow-twitch to a fast-twitch fibre transition is obtainable in mature rats.

Aging affects passive stiffness and spindle function of the rat soleus muscle

Aging affects many motor functions, notably the spinal stretch reflexes and muscle spindle sensitivity. Spindle activation also depends on the elastic properties of the structures linked to the proprioceptive receptors. We have calculated a spindle efficacy index, SEI, for old rats. This index relates the spindle sensitivity, deduced from electroneurograms recording (ENG), to the passive stiffness of the muscle. Spindle sensitivity and passive incremental stiffness were calculated during ramp and hold stretches imposed on pseudo-isolated soleus muscles of control rats (aged 4 months, n=12) and old rats (aged 24 months, n=16). SEI were calculated for the dynamic and static phases of ramp (1-80 mm/s) and for hold (0.5-2mm) stretches imposed at two reference lengths: length threshold for spindle afferents discharges, L(n) (neurogram length) and slack length, L(s). The passive incremental stiffness was calculated from the peak and steady values of passive tension, measured under the stretch conditions used for the ENG recordings, and taking into account the muscle cross-sectional area. The pseudo-isolated soleus muscles were also stretched to establish the stress-strain relationship and to calculate muscle stiffness constant. The contralateral muscle was used to count muscle spindles and spindle fibers (ATPase staining) and immunostained to identify MyHC isoforms. L(n) and L(s) lengths were not significantly different in the control group, while L(n) was significantly greater than L(s) in old muscles. Under dynamic conditions, the SEI of old muscles was the same as in controls at L(s), but it was significantly lower than in controls at L(n) due to increased passive incremental stiffness under the stretch conditions used to analyze the ENG. Under static conditions, the SEI of old muscles was significantly lower than control values at all the stretch amplitudes and threshold lengths tested, due to increased passive incremental stiffness and decreased spindle sensitivity at L(s). The muscle stiffness constant values were greater in old muscles than in controls, confirming the changes in elastic properties under passive conditions due to aging. Aging also altered the intrafusal fibers: it increased the mean number of intrafusal fibers and the contents in the slow, neonatal and developmental isoforms intrafusal of MyHC have been modified. These structural modifications do not seem great enough to counteract the loss of the spindle sensitivity or the spindle efficacy under passive conditions and after the nerve was severed. However, they may help to maintain the spindle afferent message under natural conditions and under fusimotor control.

Effects of repeated Achilles tendon vibration on triceps surae force production

Many studies reported benefits of whole-body vibration (WBV) on muscle force production. Therefore, WBV may be an important technique for muscle re-education. However vibrating platforms are heavy tools that cannot be easily used by all patients. Thus, we propose to apply vibrations directly to the Achilles tendon at rest with a portable vibrator. We investigated whether 14 days of such a vibration program would enhance triceps surae force production in healthy subjects. If successful, such a protocol could be utilized to prevent deleterious effects of hypo-activity. Twenty-nine healthy students participated in this study. The electrical evoked twitch and maximal voluntary contraction (MVC) in plantar-flexion, and electromyograms (EMG) were quantified before and at the end of the program. The vibration program consisted of 14 days of daily vibration applied at rest (duration: 1 h; frequency: 50 Hz). After the program, there was an increase in MVC associated with greater EMG of the TS. No sign of hypertrophy were found on the twitch parameters and the EMG-torque relationships. Repeated vibrations of the Achilles tendon lead to an increase in plantar-flexor activation and thus to greater force developed in voluntary conditions whilst the contractile properties assessed by the twitch are not modified. This program could be beneficial to persons with hypo-activity who are not candidates for WBV.

Motor unit recruitment and EMG power spectra during ramp contractions of a bifunctional muscle

Surface electromyograms (EMGs) were analysed on the short and long head of the biceps brachii (BBSH and BBLH) during single (F and S) or dual (F+S) flexion and supination tasks. It was confirmed, by the analysis of EMG root-mean-square (RMS) values, that the highest activations of BBSH and BBLH were obtained during a maximal dual task. This study was essentially concerned with the analysis of power spectra data obtained during progressive or ramp contractions (RCs). The shape of the power spectra established during the first second of the RCs differs between F, S and F+S tasks. Differences in mean power frequency (MPF) calculated during RCs would be representative of a recruitment of motor units (MUs) that is, at least partly, task-dependent. In order to compare MPF values calculated from RCs performed under different mechanical conditions (F, S and F+S), MPF-RMS(PSD) relationships have been established (RMS(PSD) being defined as the power spectrum density RMS). Both BBSH and BBLH exhibited initial MPF values higher in supination RC than in flexion RC. Because of plateau values reached at the same level of muscle activation whatever the task performed, the slope of the MPF-RMS(PSD) relationship was lower in S than in F. These results are in favour of MU recruitment that is, at least partly, different in F and in S conditions. Dual submaximal tasks seem to mix the activation of the F and S subpopulations of MUs as revealed by the spectral parameters obtained during F+S ramp contractions. This study could find some implication in the field of muscle rehabilitation or reinforcement.

Neuromuscular adaptations in rats trained by muscle stretch-shortening

The aim of this study was the analysis of neurophysiological, mechanical and histochemical parameters to demonstrate muscle adaptation with training. If the parameters studied were to show correlated changes, it would be possible to propose that the neural and the muscle components of motor units are both affected by the training programme used. The training consisted of repeated stretch-shortening cycles known to use extensively fast fibres. After the training period electromyographical reflex activities of the ankle plantar-flexors were recorded in awake rats and then mechanical and histochemical measurements were made on isolated soleus muscles of the control and trained rats. The reflexes studied were the H-response to electrical stimulation of the sciatic nerve and the T-response to an Achilles tendon tap. The H-response analysis indicated a decrease in reflex excitability of the trained muscles. The trained soleus muscle also presented a higher contractility as demonstrated by significantly smaller twitch contraction times and higher maximal velocities of shortening measured during tetanic contractions. The reflex and contractile muscle changes were accompanied by relative increases in the number of type II fibres. The T-response was not significantly modified by training despite the decrease in motoneuron excitability demonstrated by the decrease in H-response. This would suggest that the peripheral components of the reflex pathway such as tendon stiffness and/or spindle sensitivity might be modified by training. This would imply that both the motor and the sensory parts of a muscle are affected by training.