Unraveling intra- and intersegmental neuronal connectivity between central pattern generating networks in a multi-legged locomotor system
Animal walking results from a complex interplay of central pattern generating networks (CPGs), local sensory signals expressing position, velocity and forces generated in the legs, and coordinating signals between neighboring ones. In the stick insect, in particular, intra- and intersegmental coordination is conveyed by these sensory signals. The rhythmic activity of the CPGs, hence of the legs, can be modified by the aforementioned sensory signals. However, the precise nature of the interaction between the CPGs and these sensory signals has remained largely unknown. Experimental methods aiming at finding out details of these interactions, often apply the muscarinic acetylcholine receptor agonist, pilocarpine in order to induce rhythmic activity in the CPGs, hence in the motoneurons of the segmental ganglia. Using this general approach, we removed the influence of sensory signals and investigated the putative connections between CPGs associated with the coxa-trochanter (CTr)-joint in the different segments (legs) in more detail. The experimental data underwent phase-difference analysis and Dynamic Causal Modelling (DCM). These methods can uncover the underlying coupling structure and strength between pairs of segmental ganglia (CPGs). We set up different coupling schemes (models) for DCM and compared them using Bayesian Model Selection (BMS). Models with contralateral connections in each segment and ipsilateral connections on both sides, as well as the coupling from the meta- to the ipsilateral prothoracic ganglion were preferred by BMS to all other types of models tested. Moreover, the intrasegmental coupling strength in the mesothoracic ganglion was the strongest and most stable in all three ganglia.