To beat or not to beat: a decision taken at the network level.

The cells of the inferior olivary nucleus, the sole source of the cerebellar climbing fibers, form a network of electrically coupled neurons. Experimental observations show that these neurons produce a large repertoire of electrical signals, among which sub-threshold oscillations of the membrane potential. Simultaneous recordings from pairs of neurons and optical imaging of voltage sensitive dyes show that sub-threshold activity occurs in synchrony throughout the network. The mechanism underlying the generation of the sub-threshold oscillations is not fully understood. Experimental observations suggest that the electrical coupling is essential but insufficient for their generation. Several theoretical mechanisms have been suggested to explain these observations. Up-to-date, the most realistic model is the heterogeneity model, that assumes a certain degree of heterogeneity among olivary neurons. The heterogeneity model proposes that sub-threshold oscillations are produced by electrical coupling of neurons with the same types of ionic conductances, but with different densities. The variability in channel densities yield neurons of different functional types. The main prediction of the model is that different functional types of neurons should be found in the inferior olive. Dynamic clamp experiments support this prediction.