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Fast synaptic inhibition in the nervous system depends on the transmembrane flux of Cl -  ions based on the neuronal Cl -  driving force. Established theories regarding the determinants of Cl -  driving force have recently been questioned. Here, we present biophysical models of Cl -  homeostasis using the pump-leak model. Using numerical and novel analytic solutions, we demonstrate that the Na + /K + -ATPase, ion conductances, impermeant anions, electrodiffusion, water fluxes and cation-chloride cotransporters (CCCs) play roles in setting the Cl -  driving force. Our models, together with experimental validation, show that while impermeant anions can contribute to setting [Cl - ] i  in neurons, they have a negligible effect on the driving force for Cl -  locally and cell-wide. In contrast, we demonstrate that CCCs are well-suited for modulating Cl -  driving force and hence inhibitory signaling in neurons. Our findings reconcile recent experimental findings and provide a framework for understanding the interplay of different chloride regulatory processes in neurons.

Biophysical models reveal the relative importance of transporter proteins and impermeant anions in chloride homeostasis