A systems model for the pupil size effect

The human pupillary control system is a paradigm for linearized biological control systems. It also exhibits a series of interesting nonlinear behaviors, particularly asymmetry, “pupillary escape”, and “pupillary capture.” We present a nonlinear model in which a signal dependent upon pupil size is fed back internally to cause a change in system parameters related to gains and rates of light adaptation. The model was simulated on a digital computer, a variety of experimental data was well matched, and improvements over previous pupil models demonstrated. A candidate physiological mechanism for adaptive components of the model might have the form of an inverse “Henneman coded” neuronal pool.