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Spontaneous oscillations induced by time delays are observed in many real-world systems. Phase reduction theory for limit-cycle oscillators described by delay-differential equations (DDEs) has been developed to analyze their synchronization properties, but it is applicable only when the perturbation applied to the oscillator is sufficiently weak. In this study, we formulate a nonlinear phase-amplitude reduction theory for limit-cycle oscillators described by DDEs on the basis of the Floquet theorem, which is applicable when the oscillator is subjected to perturbations of moderate intensity. We propose a numerical method to evaluate the leading Floquet eigenvalues, eigenfunctions, and adjoint eigenfunctions necessary for the reduction and derive a set of low-dimensional nonlinear phase-amplitude equations approximately describing the oscillator dynamics. By analyzing an analytically tractable oscillator model with a cubic nonlinearity, we show that the asymptotic phase of the oscillator state in an infinite-dimensional state space can be approximately evaluated and nontrivial bistability of the oscillation amplitude caused by moderately strong periodic perturbations can be predicted on the basis of the derived phase-amplitude equations. We further analyze a model of gene-regulatory oscillator and illustrate that the reduced equations can elucidate the mechanism of its complex dynamics under nonweak perturbations, which may be relevant to real physiological phenomena such as circadian rhythm sleep disorders.

Gamma oscillations are thought to play an important role in brain function. Interneuron gamma (ING) and pyramidal interneuron gamma (PING) mechanisms have been proposed as generation mechanisms for these oscillations. However, the relation between the generation mechanisms and the dynamical properties of the gamma oscillation are still unclear. Among the dynamical properties of the gamma oscillation, the phase response function (PRF) is important because it encodes the response of the oscillation to inputs. Recently, the PRF for an inhibitory population of modified theta neurons that generate an ING rhythm was computed by the adjoint method applied to the associated Fokker-Planck equation (FPE) for the model. The modified theta model incorporates conductance-based synapses as well as the voltage and current dynamics. Here, we extended this previous work by creating an excitatory-inhibitory (E-I) network using the modified theta model and described the population dynamics with the corresponding FPE. We conducted a bifurcation analysis of the FPE to find parameter regions which generate gamma oscillations. In order to label the oscillatory parameter regions by their generation mechanisms, we defined ING- and PING-type gamma oscillation in a mathematically plausible way based on the driver of the inhibitory population. We labeled the oscillatory parameter regions by these generation mechanisms and derived PRFs via the adjoint method on the FPE in order to investigate the differences in the responses of each type of oscillation to inputs. PRFs for PING and ING mechanisms are derived and compared. We found the amplitude of the PRF for the excitatory population is larger in the PING case than in the ING case. Finally, the E-I population of the modified theta neuron enabled us to analyze the PRFs of PING-type gamma oscillation and the entrainment ability of E and I populations. We found a parameter region in which PRFs of E and I are both purely positive in the case of PING oscillations. The different entrainment abilities of E and I stimulation as governed by the respective PRFs was compared to direct simulations of finite populations of model neurons. We find that it is easier to entrain the gamma rhythm by stimulating the inhibitory population than by stimulating the excitatory population as has been found experimentally. [ABSTRACT FROM AUTHOR]