This study focuses on neuromuscular mechanisms behind ankle torque and EMG variability during a maintained isometric plantar flexion contraction. Experimentally obtained torque standard deviation (SD) and soleus, medial gastrocnemius, and lateral gastrocnemius EMG envelope mean and SD increased with mean torque for a wide range of torque levels. Computer simulations were performed on a biophysically-based neuromuscular model of the triceps surae consisting of premotoneuronal spike trains (the global input, GI) driving the motoneuron pools of the soleus, medial gastrocnemius, and lateral gastrocnemius muscles, which activate their respective muscle units. Two types of point processes were adopted to represent the statistics of the GI: Poisson and Gamma. Simulations showed a better agreement with experimental results when the GI was modeled by Gamma point processes having lower orders (higher variability) for higher target torques. At the same time, the simulations reproduced well the experimental data of EMG envelope mean and SD as a function of mean plantar flexion torque, for the three muscles. These results suggest that the experimentally found relations between torque-EMG variability as a function of mean plantar flexion torque level depend not only on the intrinsic properties of the motoneuron pools and the muscle units innervated, but also on the increasing variability of the premotoneuronal GI spike trains when their mean rates increase to command a higher plantar flexion torque level. The simulations also provided information on spike train statistics of several hundred motoneurons that compose the triceps surae, providing a wide picture of the associated mechanisms behind torque and EMG variability.

Motoneuron (MN) dendrites may be changed from a passive to an active state by increasing the levels of spinal cord neuromodulators, which activate persistent inward currents (PICs). These exert a powerful influence on MN behavior and modify the motor control both in normal and pathological conditions. Motoneuronal PICs are believed to induce nonlinear phenomena such as the genesis of extra torque and torque hysteresis in response to percutaneous electrical stimulation or tendon vibration in humans. An existing large-scale neuromuscular simulator was expanded to include MN models that have a capability to change their dynamic behaviors depending on the neuromodulation level. The simulation results indicated that the variability (standard deviation) of a maintained force depended on the level of neuromodulatory activity. A force with lower variability was obtained when the motoneuronal network was under a strong influence of PICs, suggesting a functional role in postural and precision tasks. In an additional set of simulations when PICs were active in the dendrites of the MN models, the results successfully reproduced experimental results reported from humans. Extra torque was evoked by the self-sustained discharge of spinal MNs, whereas differences in recruitment and de-recruitment levels of the MNs were the main reason behind torque and electromyogram (EMG) hysteresis. Finally, simulations were also used to study the influence of inhibitory inputs on a MN pool that was under the effect of PICs. The results showed that inhibition was of great importance in the production of a phasic force, requiring a reduced co-contraction of agonist and antagonist muscles. These results show the richness of functionally relevant behaviors that can arise from a MN pool under the action of PICs.

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Mechanically evoked reflexes have been postulated to be less sensitive to presynaptic inhibition (PSI) than the H-reflex. This has implications on investigations of spinal cord neurophysiology that are based on the T-reflex. Preceding studies have shown an enhanced effect of PSI on the H-reflex when a train of ~10 conditioning stimuli at 1 Hz was applied to the nerve of the antagonist muscle. The main questions to be addressed in the present study are if indeed T-reflexes are less sensitive to PSI and whether (and to what extent and by what possible mechanisms) the effect of low frequency conditioning, found previously for the H-reflex, can be reproduced on T-reflexes from the soleus muscle. We explored two different conditioning-to-test (C-T) intervals: 15 and 100 ms (corresponding to D1 and D2 inhibitions, respectively). Test stimuli consisted of either electrical pulses applied to the posterior tibial nerve to elicit H-reflexes or mechanical percussion to the Achilles tendon to elicit T-reflexes. The 1 Hz train of conditioning electrical stimuli delivered to the common peroneal nerve induced a stronger effect of PSI as compared to a single conditioning pulse, for both reflexes (T and H), regardless of C-T-intervals. Moreover, the conditioning train of pulses (with respect to a single conditioning pulse) was proportionally more effective for T-reflexes as compared to H-reflexes (irrespective of the C-T interval), which might be associated with the differential contingent of Ia afferents activated by mechanical and electrical test stimuli. A conceivable explanation for the enhanced PSI effect in response to a train of stimuli is the occurrence of homosynaptic depression at synapses on inhibitory interneurons interposed within the PSI pathway. The present results add to the discussion of the sensitivity of the stretch reflex pathway to PSI and its functional role.