The nodulus receives a primary vestibular afferent input from the ipsilateral labyrinth and a vestibularly related climbing-fiber input originating from the contralateral labyrinth. Previously we demonstrated that increased discharge of vestibularly evoked climbing-fiber responses (CFRs) in nodular Purkinje cells was correlated with decreased discharge of simple spikes (SSs). This left unresolved the question of whether vestibularly evoked antiphasic behavior of CFRs and SSs reflects a common neural mechanism or the activation of two separate parallel pathways. We answered this question using natural vestibular stimulation to modulate the discharge of uvula-nodular Purkinje cells recorded extracellularly in unilaterally labyrinthectomized, chloralose urethane-anesthetized rabbits. In such animals, vestibular primary afferents projecting to the uvula-nodulus as mossy fibers remained intact on the side contralateral to the unilateral labyrinthectomy. The discharge of CFRs recorded in ipsilateral nodular Purkinje cells was increased by ipsilateral roll-tilt while the discharge of SSs was increased by contralateral roll-tilt. These polarities were reversed for Purkinje cells recorded in the contralateral uvula-nodulus. The polarity of SS discharge recorded from Purkinje cells on both sides of the nodulus was opposite to that of the vestibular primary mossy-fiber afferents. SSs continued to respond to contralateral roll-tilt even when the primary vestibular afferent mossy-fiber pathway was destroyed by the unilateral labyrinthectomy. Although the discharge of SSs recorded in the contralateral uvula-nodulus was increased by contralateral roll-tilt, this modulation was reduced relative to that observed in Purkinje cells recorded in the ipsilateral uvula-nodulus. We conclude that vestibularly evoked CFRs caused the modulation of SS discharge.