The otolith vermis: A systems neuroscience theory of the Nodulus and Uvula

The Nodulus and Uvula (NU) (lobules X and IX of the cerebellar vermis) form a prominent center of vestibular information processing. Over decades, fundamental and clinical research on the NU has uncovered many aspects of its function. Those include the resolution of a sensory ambiguity inherent to inertial sensors in the inner ear, the otolith organs; the use of gravity signals to sense head rotations; and the differential processing of self-generated and externally imposed head motion. Here, I review these works in the context of a theoretical framework of information processing called the internal model hypothesis. I propose that the NU implements a forward internal model to predict the activation of the otoliths, and outputs sensory predictions errors to correct internal estimates of self-motion or to drive learning. I show that a Kalman filter based on this framework accounts for various functions of the NU, neurophysiological findings, as well as the clinical consequences of NU lesions. This highlights the role of the NU in processing information from the otoliths and supports its denomination as the "otolith" vermis.