Cortical Control of a Whisking Central Pattern Generator

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Abstract
Whether the motor cortex regulates voluntary movements by generating the motor pattern directly or by acting through subcortical central pattern generators (CPGs) remains a central question in motor control. Using the rat whisker system, an important model system of mammalian motor control, we develop an anesthetized preparation to investigate the interaction between the motor cortex and a whisking CPG. Using this model we investigate the involvement of a serotonergic component of the whisking CPG in determining whisking kinematics and the mechanisms through which drive from the CPG is converted into movements by vibrissa motor units. Consistent with an action of the vibrissa motor cortex (vMCx) on a subcortical CPG, the frequency of whisking evoked by intracortical microstimulation (ICMS) of vMCx differed significantly from the stimulation frequency, whereas whisking onset latencies correlated negatively with stimulation intensity. Further, ICMS-evoked whisking was suppressed by a serotonin receptor antagonist, supporting previous findings that the whisking CPG contains a significant serotonergic component. The amplitude of ICMS-evoked whisking was correlated with the number of active motor units—isolated from vibrissal EMGs or recorded directly from vibrissa motoneurons—and their activity level. In addition, whisking frequency was correlated with the firing rate of these motoneurons. These findings support the hypothesis that vMCx regulates whisking through its actions on a subcortical CPG.