Mechanism and function of mixed-mode oscillations in vibrissa motoneurons.

Vibrissa motoneurons in the facial nucleus innervate the intrinsic and extrinsic muscles that move the whiskers. Their intrinsic properties affect the way they process fast synaptic input from the vIRT and Bötzinger nuclei together with serotonergic neuromodulation. In response to constant current (I(app)) injection, vibrissa motoneurons may respond with mixed mode oscillations (MMOs), in which sub-threshold oscillations (STOs) are intermittently mixed with spikes. This study investigates the mechanisms involved in generating MMOs in vibrissa motoneurons and their function in motor control. It presents a conductance-based model that includes the M-type K+ conductance, g(M), the persistent Na+ conductance, g(NaP), and the cationic h conductance, g(h). For g(h) = 0 and moderate values of g(M) and g(NaP), the model neuron generates STOs, but not MMOs, in response to I(app) injection. STOs transform abruptly to tonic spiking as the current increases. In addition to STOs, MMOs are generated for g(h)>0 for larger values of I(app); the I(app) range in which MMOs appear increases linearly with g(h). In the MMOs regime, the firing rate increases with I(app) like a Devil's staircase. Stochastic noise disrupts the temporal structure of the MMOs, but for a moderate noise level, the coefficient of variation (CV) is much less than one and varies non-monotonically with I(app). Furthermore, the estimated time period between voltage peaks, based on Bernoulli process statistics, is much higher in the MMOs regime than in the tonic regime. These two phenomena do not appear when moderate noise generates MMOs without an intrinsic MMO mechanism. Therefore, and since STOs do not appear in spinal motoneurons, the analysis can be used to differentiate different MMOs mechanisms. MMO firing activity in vibrissa motoneurons suggests a scenario in which moderate periodic inputs from the vIRT and Bötzinger nuclei control whisking frequency, whereas serotonergic neuromodulation controls whisking amplitude.