Emergence and combinatorial accumulation of jittering regimes in spiking oscillators with delayed feedback

Interaction via pulses is common in many natural systems, especially neuronal. In this article we study one of the simplest possible systems with pulse interaction: a phase oscillator with delayed pulsatile feedback. When the oscillator reaches a specific state, it emits a pulse, which returns after propagating through a delay line. The impact of an incoming pulse is described by the oscillator's phase reset curve (PRC). In such a system we discover an unexpected phenomenon: for a sufficiently steep slope of the PRC, a periodic regular spiking solution bifurcates with several multipliers crossing the unit circle at the same parameter value. The number of such critical multipliers increases linearly with the delay and thus may be arbitrary large. This bifurcation is accompanied by the emergence of numerous "jittering" regimes with nonequal interspike intervals (ISIs). Each of these regimes corresponds to a periodic solution of the system with a period roughly proportional to the delay. The number of different "jittering" solutions emerging at the bifurcation point increases exponentially with the delay. We describe the combinatorial mechanism that underlies the emergence of such a variety of solutions. In particular, we show how a periodic solution exhibiting several distinct ISIs can imply the existence of multiple other solutions obtained by rearranging of these ISIs. We show that the theoretical results for phase oscillators accurately predict the behavior of an experimentally implemented electronic oscillator with pulsatile feedback.