The Goodwin model revisited: Hopf bifurcation, limit-cycle, and periodic entrainment.

The three-variable Goodwin oscillator is a minimal model demonstrating the emergence of oscillations in simple biochemical feedback systems. As a prototypical oscillator, this model was extensively studied from a theoretical point of view and applied to various biological systems, including circadian clocks. Here, we reexamine this model, derive analytically the amplitude equation near the Hopf bifurcation and investigate the effect of a periodic modulation of the oscillator. In particular, we compare the entrainment performance when the free oscillator displays either self-sustained or damped oscillations. We discuss the results in the context of circadian oscillators.