Canard induced mixed-mode oscillations in a medial entorhinal cortex layer II stellate cell model

Stellate cells (SCs) of the medial entorhinal cortex (layer II) display mixed-mode oscillatory activity, subthreshold oscillations (small-amplitude) interspersed with spikes (large amplitude), at theta frequencies (8-12 Hz). In this paper we study the mechanism of generation of such patterns in an SC biophysical (conductance-based) model. In particular, we show that the mechanism is based on the three-dimensional canard phenomenon and that the subthreshold oscillatory phenomenon is intrinsically nonlinear, involving the participation of both components (fast and slow) of a hyperpolarization-activated current in addition to the voltage and a persistent sodium current. We discuss some consequences of this mechanism for the SC intrinsic dynamics as well as for the interaction between SCs and external inhibitory inputs.