Intertrial Variability in Human Corticospinal Activity during Grasp Force Planning

Neuronal firing rate variability during planning has been found to contribute to trial-to-trial variability in primate behavior. However, in humans, whether planning related mechanisms contribute to trial-to-trial behavioral variability remains unknown. We investigated the time-course of trial-to-trial variability in corticospinal excitability (CSE) using transcranial magnetic stimulation (TMS) while subjects planned to perform a self-paced reach-to-grasp task. We hypothesized that CSE variability will be modulated during task planning and that such a modulation would explain trial-to-trial behavioral variability. Able-bodied individuals were visually cued to plan their grip force before exertion of either 30% or 5% of maximum force on an object. TMS was delivered at different time points following a cue that instructed the force level. We first modeled the relation between CSE magnitude and its variability at rest (n=12) to study the component of CSE variability during task planning that was not related to changes in CSE magnitude (n=12). We found an increase in CSE variability during task planning at 30% but not at 5% of force. This effect was temporally dissociated from the decrease in CSE magnitude. Importantly, the increase in CSE variability during planning explained 64% of inter-individual differences in time to peak force rate trial-to-trial variability. These results were found to be repeatable across studies and robust to different analysis methods. Our findings suggest that the planning-related mechanisms underlying modulation in CSE variability and CSE magnitude are distinct. Notably, the extent of modulation in planning-related variability in corticospinal system within individuals may explain their trial-to-trial behavioral variability.