Stéphane Baudry

Age-related decline in rate of torque development is accompanied by lower maximal motor unit discharge frequency during fast contractions

The aim of this study was to investigate the association between the rate of torque development and maximal motor unit discharge frequency in young and elderly adults as they performed rapid submaximal contractions with the ankle dorsiflexors. Recordings were obtained of the torque exerted by the dorsiflexors during the isometric contractions and the surface and intramuscular electromyograms (EMGs) from the tibialis anterior. The maximal rate of torque development and integrated EMG (percentage of total EMG burst) at peak rate of torque development during fast contractions were lower in elderly than young adults by 48% (P < 0.05) and 16.5% (P < 0.05), respectively. The young adults, but not the elderly adults, exhibited a positive association (r2 = 0.33; P < 0.01) between the integrated EMG computed up to the peak rate of torque development and the maximal rate of torque development achieved during the fast contractions. These age-related changes during fast voluntary contractions were accompanied by a decline (P < 0.001) in motor unit discharge frequency (19, 28, and 34% for first 3 interspike intervals, respectively) and in the percentage of units (45%; P < 0.05) that exhibited double discharges (doublets) at brief intervals (<5 ms). Because aging decreased the maximal rate of torque development of fast voluntary contractions to a greater extent ( approximately 10%) than that of an electrically evoked twitch, collectively the results indicate that the age-related decline in maximal motor unit discharge frequency likely limit, in addition to the slowing of muscle contractile properties, the performance of fast voluntary contractions.

Voluntary activation during maximal contraction with advancing age: a brief review

It is well established that the loss of muscle mass (i.e. sarcopenia) is the primary factor contributing to the reduction in muscle force with ageing. Based on the observation that force declines at a faster rate than muscle mass, neural alterations are also thought to contribute to muscle weakness by reducing central drive to the agonist muscles and by increasing coactivation of the antagonist muscles. Researchers have attempted to quantify the contribution of impaired voluntary drive to the decline in muscle force using superimposed electrical stimulation during maximal voluntary contractions (MVCs) and by recording surface electromyographic (EMG) activity. Although reduced voluntary activation of agonist muscles and increased coactivation of antagonist muscles during a MVC have been reported with advancing age, such changes are not supported by all studies. These discrepancies may be explained by differences in sensitivity between the methods used to assess voluntary activation, as well as differences between the characteristics of the study population, the muscle group that is tested, and the type of contraction that is performed. The objective of this review is to summarize current knowledge regarding the activation of agonist and antagonist muscles during MVC in elderly and to try to clarify the disparities in literature concerning the influence of a possible deficit in voluntary activation on the maximal force capacity of muscles in elderly adults.

Unraveling the neurophysiology of muscle fatigue

Despite 100years of research since the seminal work of Angelo Mosso (1846-1910), our understanding of the interactions between the nervous system and muscle during the performance of fatiguing contractions remains rather rudimentary. Although the nervous system simply needs to provide an activation signal that will elicit the net muscle torque required for a prescribed action, changes in the number and diversity of synaptic inputs that must be integrated by the spinal motor neurons to accommodate the changes in the force-producing capabilities of the muscle fibers complicate the process of generating the requisite activation signal. This brief review examines two ways in which the activation signal can be compromised during sustained contractions and thereby contribute to the rate at which the muscles fatigue. These examples provide insight on the types of adjustments that occur in the nervous system during fatiguing contractions, but emphasize that much remains to be learned about the physiological processes that contribute to the phenomenon known as muscle fatigue.

Aging causes a reorganization of cortical and spinal control of posture

Classical studies in animal preparations suggest a strong role for spinal control of posture. In humans it is now established that the cerebral cortex contributes to postural control of unperturbed and perturbed standing. The age-related degeneration and accompanying functional changes in the brain, reported so far mainly in conjunction with simple manual motor tasks, may also affect the mechanisms that control complex motor tasks involving posture. This review outlines the age-related structural and functional changes at spinal and cortical levels and provides a mechanistic analysis of how such changes may be linked to the behaviorally manifest postural deficits in old adults. The emerging picture is that the age-related reorganization in motor control during voluntary tasks, characterized by differential modulation of spinal reflexes, greater cortical activation and cortical disinhibition, is also present during postural tasks. We discuss the possibility that this reorganization underlies the increased coactivation and dual task interference reported in elderly. Finally, we propose a model for future studies to unravel the structure-function-behavior relations in postural control and aging.

Postactivation potentiation in human muscle is not related to the type of maximal conditioning contraction.

The mechanical performance of a muscle can be enhanced by preceding contractile activity, such as occurs with postactivation potentiation. To investigate whether the type of contraction influences the extent of potentiation, the effects of 6‐s maximal isometric (ISO), concentric (CON), and eccentric (ECC) maximal voluntary contractions (MVC) on the muscle twitch were compared in the tibialis anterior of nine subjects. The study also examined the effect of postactivation potentiation on the force evoked by the second (C2) and third (C3) responses of two‐pulse (PT2) and three‐pulse (PT3) trains that were delivered at a 10‐ms interpulse interval. The results showed that immediately after the conditioning MVC, twitch torque (Pt) and its maximal rate of torque development (+dPt/dt) and relaxation (‐dPt/dt) were significantly enhanced, without any change in contraction time (CT), half‐relaxation time (½RT), and compound muscle action potential (M wave). The extent of Pt potentiation was similar for all MVC modalities, and the mean maximal values ranged from 150% to 180%. Furthermore, postactivation potentiation was greater for the single pulse compared with PT2 and PT3 responses. All parameters returned to initial values within 7–10 min. Although Pt (or C1) was potentiated more than was C2 and C3, its decline over time was proportionally more rapid than those for C2 and C3. We conclude that postactivation potentiation was not related to the type of conditioning MVC under these experimental conditions. The observation that postactivation potentiation increased C1 more than C2 and C3 indicates that a saturation process limits the extent of potentiation during the summation of successive responses to a train of stimuli. These results have practical application in the design of functional electrical stimulation protocols. Muscle Nerve, 2004

Presynaptic Modulation of Ia Afferents in Young and Old Adults When Performing Force and Position Control

The present work investigated presynaptic modulation of Ia afferents in the extensor carpi radialis (ECR) when young and old adults exerted a wrist extension force either to support an inertial load (position control) or to achieve an equivalent constant torque against a rigid restraint (force control) at 5, 10, and 15% of the maximal force. H reflexes were evoked in the ECR by stimulating the radial nerve above the elbow. A conditioning stimulus was applied to the median nerve above the elbow to assess presynaptic inhibition of homonymous Ia afferents (D1 inhibition) or at the wrist (palmar branch) to assess the ongoing presynaptic inhibition of heteronymous Ia afferents that converge onto the ECR motor neuron pool (heteronymous Ia facilitation). The young adults had less D1 inhibition and greater heteronymous Ia facilitation during the position task (79 and 132.1%, respectively) compared with the force task (69.1 and 115.1%, respectively, P < 0.05). In contrast, the old adults exhibited no difference between the two tasks for either D1 inhibition ( approximately 72%) or heteronymous Ia facilitation ( approximately 114%). Contraction intensity did not influence the amount of D1 inhibition or heteronymous Ia facilitation for either group of subjects. The amount of antagonist coactivation was similar between tasks for young adults, whereas it was greater in the position task for old adults (P = 0.02). These data indicate that in contrast to young adults, old adults did not modulate presynaptic inhibition of Ia afferents when controlling the position of a compliant load but rather increased coactivation of the antagonist muscle.

Age‐related influence of vision and proprioception on Ia presynaptic inhibition in soleus muscle during upright stance

•  The observed decrease in balance with ageing, vision suppression and compliance of the support surface may involve differential modulation of Ia afferent feedback from leg muscles. •  The modulation of Ia presynaptic inhibition for the soleus muscle was assessed in young and elderly adults when standing in normal and modified visual (eyes closed) and proprioceptive conditions (foam support). •  The results suggest that presynaptic inhibition of Ia afferents increased when vision was suppressed and when standing on a foam mat, but more so in elderly adults, for whom the increase in Ia presynaptic inhibition was associated with greater activity of the leg muscles. •  Young and elderly adults appear to rely less on segmental muscle afferent feedback to control the activation of leg muscles during upright stance when vision and proprioception are altered.

Load type influences motor unit recruitment in biceps brachii during a sustained contraction

Twenty subjects participated in four experiments designed to compare time to task failure and motor-unit recruitment threshold during contractions sustained at 15% of maximum as the elbow flexor muscles either supported an inertial load (position task) or exerted an equivalent constant torque against a rigid restraint (force task). Subcutaneous branched bipolar electrodes were used to record single motor unit activity from the biceps brachii muscle during ramp contractions performed before and at 50 and 90% of the time to failure for the position task during both fatiguing contractions. The time to task failure was briefer for the position task than for the force task (P=0.0002). Thirty and 29 motor units were isolated during the force and position tasks, respectively. The recruitment threshold declined by 48 and 30% (P=0.0001) during the position task for motor units with an initial recruitment threshold below and above the target force, respectively, whereas no significant change in recruitment threshold was observed during the force task. Changes in recruitment threshold were associated with a decrease in the mean discharge rate (-16%), an increase in discharge rate variability (+40%), and a prolongation of the first two interspike intervals (+29 and +13%). These data indicate that there were faster changes in motor unit recruitment and rate coding during the position task than the force task despite a similar net muscle torque during both tasks. Moreover, the results suggest that the differential synaptic input observed during the position task influences most of the motor unit pool.

Insights into the neural control of eccentric contractions

The purpose of this brief review is to examine our current knowledge of the neural control of eccentric contractions. The review focuses on three main issues. The first issue considers the ability of individuals to activate muscles maximally during eccentric contractions. Most studies indicate that, regardless of the experimental approach (surface EMG amplitude, twitch superimposition, and motor unit recordings), it is usually more difficult to achieve full activation of a muscle by voluntary command during eccentric contractions than during concentric and isometric contractions. The second issue is related to the specificity of the control strategy used by the central nervous system during submaximal eccentric contractions. This part underscores that although the central nervous system appears to employ a single size-related strategy to activate motoneurons during the different types of contractions, the discharge rate of motor units is less during eccentric contractions across different loading conditions. The last issue addresses the mechanisms that produce this specific neural activation. This section indicates that neural adjustments at both supraspinal and spinal levels contribute to the specific modulation of voluntary activation during eccentric contractions. Although the available information on the control of eccentric contractions has increased during the last two decades, this review indicates that the exact mechanisms underlying the unique neural modulation observed in this type of contraction at spinal and supraspinal levels remains unknown and their understanding represents, therefore, a major challenge for future research on this topic.

Age-related changes in the behavior of the muscle-tendon unit of the gastrocnemius medialis during upright stance

Mechanical properties of the muscle-tendon unit change with aging, but it is not known how these modifications influence the control of lower leg muscles during upright stance. In this study, young and elderly adults stood upright on a force platform with and without vision while muscle architecture and myotendinous junction movements (expressed relative to the change in the moment on the x-axis of the force platform) were recorded by ultrasonography and muscle activity by electromyography. The results show that the maximal amplitude of the sway in the antero-posterior direction was greater in elderly adults (age effect, P < 0.05) and was accompanied by an increase in lower leg muscle activity compared with young adults. Moreover, the data highlight that fascicles shorten during forward sway and lengthen during backward sways but more so for young (-4 ± 3 and -4 ± 3 mm/Nm, respectively) than elderly adults (-0.7 ± 3 and 0.8 ± 3 mm/Nm, respectively; age × sway, P < 0.001). Concurrently, the pennation angle increased and decreased during forward and backward sways, respectively, with greater changes in young than elderly adults (age × sway, P < 0.001). In contrast, no significant differences were observed between age groups for tendon lengthening and shortening during sways. The results indicate that, compared with young, elderly adults increase the stiffness of the muscular portion of the muscle-tendon unit during upright stance that may compensate for the age-related decrease in tendon stiffness. These observations suggest a shift in the control strategy used to maintain balance.