Regulation of the Crab Heartbeat by Crustacean Cardioactive Peptide (CCAP): Central and Peripheral Actions

Abstract

In regulating neurophysiological systems, neuromodulators exert multiple actions at multiple sites in such a way as to control the activity in an integrated manner. We are studying how this happens in a simple central pattern generator (CPG)–effector system, the heart of the blue crab Callinectes sapidus. The rhythmic contractions of this heart are neurogenic, driven by rhythmic motor patterns generated by the cardiac ganglion (CG). In this study, we used anatomical and physiological methods to examine the sources and actions on the system of crustacean cardioactive peptide (CCAP). Immunohistochemical localization revealed a plexus of CCAP-immunoreactive fibers in the pericardial organs (POs), neurohemal structures from which blood-borne neurohormones reach the heart. Combined backfill and immunohistochemical experiments indicated that the CCAP in the POs originated from a large contralateral neuron in each thoracic neuromere. In physiological experiments, we examined the actions of exogenous CCAP on the intact working heart, on the semi-intact heart in which we could record the motor patterns as well as the muscle contractions, and on the isolated CG. CCAP had strong positive inotropic and chronotropic effects. Dissection of these effects in terms of dose dependency, time course, and the preparation type in which they occurred suggested that they were produced by the interaction of three primary actions of CCAP exerted both on the heart muscle and on the CG. We conclude that CCAP released from the POs as a neurohormone regulates the crab heart by multiple actions on both the central and peripheral components of this model CPG–effector system.