Mixed (chemical and electrical) synapses on frog spinal motoneurons

Freeze-fracture replicas and ultrathin sections were used to characterize the gap junctions on the somata and large dendrites of frog motoneurons found earlier by Sonnhof et al. (1977). In freeze-fracture replicas one of the specific features of these relatively frequent gap junctions is the presence of circular regions of non-junctional membrane ("fenestrae") within areas of typical gap junction appearance displaying P-face particles or E-face pits. Such "fenestrated" gap junctions are mostly associated with membrane specializations indicative of the active zone of a chemical synapse (including vesicle attachment sites in non anaesthetized animals) to constitute mixed synapses. These findings could be verified in ultrathin sections, which revealed that the vesicles of the chemical component of the mixed synapses were spherical and agranular. Our results suggest that the mixed synapses are predominantly axo-somatic and axo-dendritic. The existence of dendro-dendritic gap junctions in the ventral horn region as described by Sotelo and Taxi (1970) was verified in ultrathin sections; they were rare, solely electrotonic in character, and probably represent the morphological basis for the VR-EPSP (Katz and Miledi, 1963; Kubota and Brookhart, 1963), i.e. electrotonic coupling between motoneurons of different spinal segments (Washizu, 1960). Electrotonic coupling can also be demonstrated between motoneurons and afferent fibers of the dorsal root and the lateral column. Electrotonic potentials recorded within motoneurons during electrical stimulation of dorsal root or lateral column precede the chemical postsynaptic potentials; after Mn2+-blockade of chemical synaptic transmission, the electrotonic component persists. Some fibers of these afferent pathways are therefore assumed to act monosynaptic on the motoneuron via mixed axo-somatic and axo-dendritic synapses.