Nonlinear behavior of sinusoidally forced pyloric pacemaker neurons.

Periodic current forcing was used to investigate the intrinsic dynamics of a small group of electrically coupled neurons in the pyloric central pattern generator (CPG) of the lobster. This group contains three neurons, namely the two pyloric dilator (PD) motoneurons and the anterior burster (AB) interneuron. Intracellular current injection, using sinusoidal waveforms of varying amplitude and frequency, was applied in three configurations of the pacemaker neurons: 1) the complete pacemaker group, 2) the two PDs without the AB, and 3) the AB neuron isolated from the PDs. Depending on the frequency and amplitude of the injected current, the intact pacemaker group exhibited a wide variety of nonlinear behaviors, including synchronization to the forcing, quasiperiodicity, and complex dynamics. In contrast, a single, broad 1:1 entrainment zone characterized the response of the PD neurons when isolated from the main pacemaker neuron AB. The isolated AB responded to periodic forcing in a manner similar to the complete pacemaker group, but with wider zones of synchronization. We have built an analog electronic circuit as an implementation of a modified Hindmarsh-Rose model for simulating the membrane potential activity of pyloric neurons. We subjected this electronic model neuron to the same periodic forcing as used in the biological experiments. This four-dimensional electronic model neuron reproduced the autonomous oscillatory firing patterns of biological pyloric pacemaker neurons, and it expressed the same stationary nonlinear responses to periodic forcing as its biological counterparts. This adds to our confidence in the model. These results strongly support the idea that the intact pyloric pacemaker group acts as a uniform low-dimensional deterministic nonlinear oscillator, and the regular pyloric oscillation is the outcome of cooperative behavior of strongly coupled neurons, having different dynamical and biophysical properties when isolated.