Modulation of Circuit Feedback Specifies Motor Circuit Output

Bidirectional communication (i.e., feedforward and feedback pathways) between functional levels is common in neural systems, but in most systems little is known regarding the function and modifiability of the feedback pathway. We are exploring this issue in the crab (Cancer borealis) stomatogastric nervous system by examining bidirectional communication between projection neurons and their target central pattern generator (CPG) circuit neurons. Specifically, we addressed the question of whether the peptidergic post-oesophageal commissure (POC) neurons trigger a specific gastric mill (chewing) motor pattern in the stomatogastric ganglion solely by activating projection neurons, or by additionally altering the strength of CPG feedback to these projection neurons. The POC-triggered gastric mill rhythm is shaped by feedback inhibition onto projection neurons from a CPG neuron. Here, we establish that POC stimulation triggers a long-lasting enhancement of feedback-mediated IPSC/Ps in the projection neurons, which persists for the same duration as POC-gastric mill rhythms. This strengthened CPG feedback appears to result from presynaptic modulation, because it also occurs in other projection neurons whose activity does not change after POC stimulation. To determine the function of this strengthened feedback synapse, we compared the influence of dynamic-clamp-injected feedback IPSPs of pre- and post-POC amplitude into a pivotal projection neuron after POC stimulation. Only the post-POC amplitude IPSPs elicited the POC-triggered activity pattern in this projection neuron and enabled full expression of the POC-gastric mill rhythm. Thus, the strength of CPG feedback to projection neurons is modifiable and can be instrumental to motor pattern selection.