A functional asymmetry in the Leech Heartbeat Timing Network is revealed by driving the network across various cycle periods

We tested predictions of a computational model (Hill et al., 2002) of the leech heartbeat timing network. The timing network consists of two segmental oscillators located in the third (G3) and fourth (G4) segmental ganglia. Each oscillator consists of two reciprocally inhibitory oscillator interneurons along with the coordinating interneuron fibers that link them. In the model, the network was driven to cycle periods around the normal period of the network by repeatedly stimulating one of the paired oscillator interneurons in G3 or G4. Here we replicate these experiments in the biological system. The model predicts that the G3 and G4 oscillators can entrain the timing network to periods faster but not slower than the inherent period of the nondriven ("follower") oscillator and that they can do so symmetrically. The biological system can be driven to periods both faster (such that the driven oscillator leads in phase) and slower (such that the driven oscillator lags in phase) than the inherent period of the timing network. Although both oscillators can entrain the network, the G4 oscillator does so over a narrower range of periods. Two differences between the assumptions of the model and the properties of the biological network, spike frequency adaptation in coordinating interneurons and asymmetry in the connections from the oscillator interneurons to the coordinating interneurons, may account for these discrepancies. Individual coordinating interneurons were also able to entrain the oscillators but with little effect of the phase relationship between the oscillators, suggesting that phase relations are determined by properties inherent to the oscillator interneurons.