The role of behavioral diversity in determining the extent to which the cardiac ganglion is modulated in three species of crab

Central pattern generators (CPGs) are neural networks that generate the rhythmic outputs that control behaviors such as locomotion, respiration, and chewing. The stomatogastric nervous system (STNS), which contains the CPGs that control foregut movement, and the cardiac ganglion (CG), which is a CPG that controls heartbeat, are two commonly studied systems in decapod crustaceans. Neuromodulators are locally or hormonally released neuropeptides and amines that change the output patterns of CPGs like the STNS and CG to allow behavioral flexibility. We have hypothesized that neuromodulation provides a substrate for the evolution of behavioral flexibility, and as a result, systems exhibiting more behavioral flexibility are modulated to a greater degree. To examine this hypothesis, we evaluated the extent to which the STNS and the CG are modulated in the majoid crab species Chionoecetes opilio, Libinia emarginata, and Pugettia producta. C. opilio and L. emarginata are opportunistic feeders, whereas P. producta has a highly specialized kelp diet. We predicted that opportunistic feeding crabs that chew and process a wide variety of food types would exhibit greater STNS neuromodulatory capacity than those with a specialized diet. The STNS of L. emarginata and C. opilio responded to the seven endogenous neuromodulators oxotremorine, dopamine, CabTrp Ia, CCAP, myosuppressin, proctolin, and RPCH, whereas the STNS of P. producta only responded to proctolin, oxotremorine, myosuppressin, RPCH (25% of the time), variably to dopamine, and not at all to CabTrp and CCAP. Because P. producta, L. emarginata, and C. opilio all belong to the Majoidea superfamily, their primary distinctions are their feeding habits. For this reason, we further predicted that there would be no relationship between diet and modulatory capacity in the cardiac ganglion (CG) of the neurogenic heart. This would suggest that a lack of STNS modulatory capacity in P. producta relative to L. emarginata and C. opilio is specific to evolved foregut function. Whole-heart recordings from P. producta indicated that, unlike the STNS, the CG responds to CabTrp and CCAP. P. producta hearts also responded to oxotremorine and inconsistently to dopamine and proctolin. The CG of C. opilio was modulated by CabTrp, CCAP, dopamine, proctolin, myosuppressin, and oxotremorine, but not RPCH. The CG of L. emarginata responded to CCAP, and inconsistently to CabTrp, dopamine, and proctolin, but not to myosuppressin, RPCH, and surprisingly oxotremorine. Although cardiac responses were not identical between species, opportunistic and specialist feeders responded more similarly to the modulators tested in the heart than in the STNS. Notably, P. producta responded to each modulator in a similar manner to C. opilio and/or L. emarginata. However, L. emarginata’s surprising lack of cardiac response to oxotremorine suggests that phylogenetic closeness may not control for differences in CG and STNS function between species. Nevertheless, sample sizes of all three species were quite small, and individual differences lead to inconsistencies in the data. As a result, sample size must be enlarged to draw firm conclusions.