Mixed-mode bursting oscillations: Dynamics created by a slow passage through spike-adding canard explosion in a square-wave burster

This article concerns the phenomenon of Mixed-Mode Bursting Oscillations (MMBOs). These are solutions of fast-slow systems of ordinary differential equations that exhibit both small-amplitude oscillations (SAOs) and bursts consisting of one or multiple large-amplitude oscillations (LAOs). The name MMBO is given in analogy to Mixed-Mode Oscillations, which consist of alternating SAOs and LAOs, without the LAOs being organized into burst events. In this article, we show how MMBOs are created naturally in systems that have a spike-adding bifurcation or spike-adding mechanism, and in which the dynamics of one (or more) of the slow variables causes the system to pass slowly through that bifurcation. Canards are central to the dynamics of MMBOs, and their role in shaping the MMBOs is two-fold: saddle-type canards are involved in the spike-adding mechanism of the underlying burster and permit one to understand the number of LAOs in each burst event, and folded-node canards arise due to the slow passage effect and control the number of SAOs. The analysis is carried out for a prototypical fourth-order system of this type, which consists of the third-order Hindmarsh-Rose system, known to have the spike-adding mechanism, and in which one of the key bifurcation parameters also varies slowly. We also include a discussion of the MMBO phenomenon for the Morris-Lecar-Terman system. Finally, we discuss the role of the MMBOs to a biological modeling of secreting neurons.