Insight into the transfer function,gain, and oscillation onset for the pupil light reflex using nonlinear delay-differential equations

Analogies are drawn between a physiologically relevant nonlinear delay-differential equation (DDE) model for the pupil light reflex and servo control analytic approaches. This DDE is shown to be consistent with the measured open loop transfer function and hence physiological insight can be obtained into the gain of the reflex and its properties. A Hopf bifurcation analysis of the DDE shows that a limit cycle oscillation in pupil area occurs when the first mode of the characteristic equation becomes unstable. Its period agrees well with experimental measurements. Beyond the point of instability onset, more modes become unstable corresponding to multiple encirclings of (-1, 0) on the Nyquist plot. These modes primarily influence the shape of the oscillation. Techniques from dynamical systems theory, e.g. bifurcation analysis, can augment servo control analytic methods for the study of oscillations produced by nonlinear neural feedback mechanisms.