LETTER Communicated by Michael DeWeese Dynamics of Nonlinear Feedback Control

Feedback control in neural systems is ubiquitous. Here we study the mathematics of nonlinear feedback control.We comparemodels inwhich the input is multiplied by a dynamic gain (multiplicative control) with models in which the input is divided by a dynamic attenuation (divi-sive control). The gain signal (resp. the attenuation signal) is obtained through a concatenation of an instantaneous nonlinearity and a linear low-pass filter operating on the output of the feedback loop. For input steps, the dynamics of gain and attenuation can be very different, de-pending on the mathematical form of the nonlinearity and the ordering of the nonlinearity and the filtering in the feedback loop. Further, the dy-namics of feedback control can be strongly asymmetrical for increment versus decrement steps of the input. Nevertheless, for each of the mod-els studied, the nonlinearity in the feedback loop can be chosen such that immediately after an input step, the dynamics of feedback control is symmetric with respect to increments versus decrements. Finally, we study the dynamics of the output of the control loops and find conditions under which overshoots and undershoots of the output relative to the steady-state output occur when the models are stimulated with low-pass filtered steps. For small steps at the input, overshoots and undershoots of the output do not occur when the filtering in the control path is faster than the low-pass filtering at the input. For large steps at the input, how-ever, results depend on the model, and for some of the models, multiple overshoots and undershoots can occur even with a fast control path.