Beta inputs to motor neurons do not directly contribute to volitional force modulation.

Neural oscillatory activity in the beta band (13-30 Hz) is prominent in the brain and it is transmitted partly linearly to the spinal cord and muscles. Multiple views on the functional relevance of beta activity in the motor system have been proposed. Previous simulation work suggested that pools of spinal motoneurons (MNs) receiving a common beta input could demodulate this activity, transforming it into low-frequency neural drive that could alter force production in muscles. This may suggest that common beta inputs to muscles have a direct role in force modulation. Here we report the experimental average levels and ranges of common beta activity in spinal MNs projecting to single muscles and use a computational model of a MN pool to test if the experimentally observed beta levels in MNs can influence force. When beta was modelled as a continuous activity, the amplitude needed to produce non-negligible changes in force corresponded to beta representation in the MN pool that was far above the experimental observations. On the other hand, when beta activity was modelled as short-lived events (i.e. bursts of beta activity separated by intervals without beta oscillations), this activity approximated levels that could cause small changes in force with estimated average common beta inputs to the MNs compatible with the experimental observations. Nonetheless, bursting beta is unlikely to be used for force control due to the temporal sparsity of this activity. It is therefore concluded that beta oscillations are unlikely to contribute to the voluntary modulation of force. KEY POINTS: It has been previously proposed that beta (13-30 Hz) common inputs to a motor neuron pool may have a non-linear effect in voluntary force control. The needed strength of beta oscillations to modulate forces has not been analysed yet. Based on computer simulations, we show that sustained beta inputs to a spinal motoneuron pool at physiologically reported levels have minimal effect on force. Levels of sustained beta rhythmic activity that can cause a significant change in force are not compatible with experimental observations of intramuscular coherence in human skeletal muscles.

Standard intensities of transcranial alternating current stimulation over the motor cortex do not entrain corticospinal inputs to motor neurons.

Transcranial alternating current stimulation (TACS) is commonly used to synchronize a cortical area and its outputs to the stimulus waveform, but gathering evidence for this based on brain recordings in humans is challenging. The corticospinal tract transmits beta oscillations (∼21 Hz) from the motor cortex to tonically contracted limb muscles linearly. Therefore, muscle activity may be used to measure the level of beta entrainment in the corticospinal tract due to TACS over the motor cortex. Here, we assessed whether TACS is able to modulate the neural inputs to muscles, which would provide indirect evidence for TACS-driven neural entrainment. In the first part of the study, we ran simulations of motor neuron (MN) pools receiving inputs from corticospinal neurons with different levels of beta entrainment. Results suggest that MNs are highly sensitive to changes in corticospinal beta activity. Then, we ran experiments on healthy human subjects (N = 10) in which TACS (at 1 mA) was delivered over the motor cortex at 21 Hz (beta stimulation), or at 7 Hz or 40 Hz (control conditions) while the abductor digiti minimi or the tibialis anterior muscle were tonically contracted. Muscle activity was measured using high-density electromyography, which allowed us to decompose the activity of pools of motor units innervating the muscles. By analysing motor unit pool activity, we observed that none of the TACS conditions could consistently alter the spectral contents of the common neural inputs received by the muscles. These results suggest that 1 mA TACS over the motor cortex given at beta frequencies does not entrain corticospinal activity. KEY POINTS: Transcranial alternating current stimulation (TACS) is commonly used to entrain the communication between brain regions. It is challenging to find direct evidence supporting TACS-driven neural entrainment due to the technical difficulties in recording brain activity during stimulation. Computational simulations of motor neuron pools receiving common inputs in the beta (∼21 Hz) band indicate that motor neurons are highly sensitive to corticospinal beta entrainment. Motor unit activity from human muscles does not support TACS-driven corticospinal entrainment.