Synaptic organization in the pacemaker nucleus of a medium-frequency weakly electric fish, Eigenmannia sp

The medullary command nucleus (MCN) of the medium-frequency weakly electric fish, Eigenmannia sp., contains two types of neurones, namely large and small cells, which are embedded in a neuropile of large and small myelinated fibers. Using serial semi-thin and ultra-thin sectioning, combined with HRP labelling it has been established that both cell types possess rich dendritic arborization and large myelinated axons. Only the axons of the large cells leave the nucleus and these constitute the unique output of the MCN. Axon branching has been observed only in the axons of small cells and their collaterals show an exclusively intranuclear course. Two types of synaptic terminals have been found on large as well as on small cells: (1) large club endings forming both gap (electrotonic) junctions and polarized chemical synapses, which often appear at the same junction constituting morphologically mixed synapses; and (2) small bouton-like terminals forming exclusively chemical synaptic contacts. No differences between the two neuron types could be detected with respect to the arrangement of the synaptic contacts: club endings and small bouton-like terminals synapse on dendritic processes as well as on perikarya, while the unmyelinated initial segments were always found to be free of synaptic contacts. Large and small cells were found to be simultaneously connected by the same club ending or small bouton-like terminal: in the case of club endings by means of gap junctions and chemical synapses, whereas in the case of boutons by chemical synapses only. Club endings sometimes form gap junctions with each other. The possible role of these unusual synaptic connections in local synchronization is suggested. Club endings originate from the large axons of small cells, while small bouton-like terminals originate from the fine myelinated fibers of extranuclear origin. In Eigenmannia, small cells, being connected to large cells as well as to each other by axo-somatic and axodendritic synapses, can be considered as the pacemaker cells of the MCN whereas large cells are relay cells. Small bouton-like terminals may convey exogeneous impulses towards the MCN exerting modulatory effects at both pacemaker and relay cell levels. The greater variety of ultrastructural correlates established in the MCN of Eigenmannia, in comparison with Sternarchus (see also ref. 16), suggests increased modulation possibilities in the former fish's EOD behaviour.