Striatal pathways dissociably control action counting and goal-directed steering

The basal ganglia (BG) are central to voluntary action, yet how they organize complex behavior remains unclear. Using a novel operant counting task, we trained mice to perform a specific number of lever presses to obtain a reward, enabling quantification of continuous kinematics and discrete actions. Stimulation of direct pathway and indirect pathway neurons (dSPNs and iSPNs) exert bidirectional and dissociable influences on both movement steering and press count: activation of dSPNs steers mice contraversively and extends press sequences, whereas activation of iSPNs steers mice ipsiversively and prematurely terminates press sequences. Calcium imaging reveals dSPNs and iSPNs that tracked physical approach or count progress, with ramping activity patterns consistent with accumulation and discharge dynamics. The difference between dSPN and iSPN population activity scales with proximity to spatial and numerical goals. These findings show that the BG implement a push-pull controller to integrate kinematics and action counting to steer progress toward goals.