Transfer of the heart pacemaker during juvenile development in the isopod crustacean Ligia exotica

Developmental changes in heartbeat pacemaker mechanisms were examined electrophysiologically in the isopod crustacean Ligia exotica. The heartbeat of embryos and early juveniles was myogenic. The heart muscle cells were coupled electrically, and no localized pacemaker activity was found in the heart. In newly hatched juveniles, the cardiac ganglion exhibited no spontaneous activity, although stimulation of the cardiac ganglion produced excitatory junctional potentials (EJPs) in the heart muscle. The myogenic activity of the heart was reset and entrained by the EJPs evoked by ganglionic stimulation. During juvenile development, spontaneous EJPs appeared irregularly in the heart muscle. Later in development, the cardiac ganglion started rhythmic bursting, and each muscle response followed a ganglionic burst discharge and overlapped the EJPs evoked by ganglionic activity. At this point, the activity of the cardiac ganglion was suppressed by application of tetrodotoxin (TTX); however, even in old adults, both muscle activity and the heartbeat continued following TTX application. Heartbeat frequency was lower in TTX-containing saline than in normal saline. These results show that, during juvenile development, the heart pacemaker is transferred from the heart muscle to the cardiac ganglion, which becomes the primary pacemaker and entrains the heart muscle activity to a higher frequency via EJPs.