| Bifurcations and complex oscillations in the human pupil light reflex (PLR) are studied. Autonomous pupil area oscillations are produced by substituting electronically controllable nonlinear feedback for the normal negative feedback of this reflex. A physiologically sound theoretical framework in which to study pupillary oscillations is developed. The model, framed as a delay-differential equation (DDE), agrees quantitatively with the simpler periodic behaviors and qualitatively with the complex behaviors. Much of the aperiodicity in the data can be ascribed to noise and transients rather than to chaos. The critical behavior of the PLR at oscillation onset is different with piecewise constant rather than smooth negative feedback. In the former, relative fluctuations in period are larger than those in amplitude, and vice versa in the latter. Properties of the time solutions and densities of a stochastic DDE are used to explain this experimental result. The Hopf bifurcation in this system is postponed by both additive and multiplicative colored noise. Theoretical insight into the behavior of stationary densities of DDE's and the origin of the postponement is given, and implications for analyzing bifurcations in neural delayed feedback systems are discussed. |
