Equilibrium States and Their Stability in the Head-Direction Ring Network.

Head-direction cells have been found in several areas in the mammalian brains. The firing rate of an ideal head-direction cell reaches its peak value only when the animal's head points in a specific direction, and this preferred direction stays the same regardless of spatial location. In this paper we combine mathematical analytical techniques and numerical simulations to fully analyze the equilibrium states of a generic ring attractor network, which is a widely used modeling framework for the head-direction system. Under specific conditions, all solutions of the ring network are bounded, and there exists a Lyapunov function that guarantees the stability of the network for any given inputs, which may come from multiple sources in the biological system, including self-motion information for inertially based updating and landmark information for calibration. We focus on the first few terms of the Fourier series of the ring network to explicitly solve for all possible equilibrium states, followed by a stability analysis based on small perturbations. In particular, these equilibrium states include the standard single-peaked activity pattern as well as double-peaked activity pattern, whose existence is unknown but has testable experimental implications. To our surprise, we have also found an asymmetric equilibrium activity profile even when the network connectivity is strictly symmetric. Finally we examine how these different equilibrium solutions depend on the network parameters and obtain the phase diagrams in the parameter space of the ring network.