Modulation of pacemaker activity by IPSP and brief length perturbations in the crayfish stretch receptor

We have studied the influences of brief stretches and releases and of inhibitory postsynaptic potentials (IPSPs) on pacemaker activity of the crayfish stretch receptor (RM1). Stimuli shift or reset the ongoing rhythm. Resettings were different if evaluated in interspike intervals containing perturbations, or in succeeding ones, and are referred to as early and late, respectively. Early resetting revealed that stretches and releases or IPSPs advance and delay, respectively, the next spike. With small stretches and releases or IPSPs, effects depend on the timing of the perturbation relative to the previous spike or phase, but above a characteristic mechanical perturbation amplitude the next spike fires at a fixed latency, invariant with the phase. Of particular interest was the finding that during late resetting the first successive intervals following stretches and releases or IPSPs were longer and shorter, respectively, than the period. This led, in approximately 50% of the cases, to a gradual recovery of the original pacemaker beat in the sense that spikes fire timed as if the early rhythm shift had not occurred. In conclusion, the recent firing history is essential in determining the RM1's response. The receptor's sensitivity is a complex nonlinear and periodic function of the pacemaker activity, and the response is due to interactions between pacemaker- and perturbation-induced transmembrane ionic currents. Although several alternative mechanisms may underly beat recovery, the results suggest that at least two coupled oscillators, one perturbable and the other not, provide a better explanation than a single oscillator. The physiological significance of resettings is unknown, but the early rhythm shift may synchronize RM1s in several segments when the animal's tail is moved, and conversely recovery would reduce synchrony, with obvious influences on shared postsynaptic neurons.