Excitatory input to burst neurons from the labyrinth and its mediating pathway in the cat: location and functional characteristics of burster-driving neurons

Summary 1. Spikes of single neurons were recorded extracellularly in the cat prepositus hypoglossi nucleus and the underlying reticular formation and were identified as type II neurons by horizontal rotation. Among these neurons, those activated by contralateral vestibular nerve stimulation with short latencies (1.5–3.0 ms) were selected for further study. 2. A class of these identified neurons was antidromically activated from the contralateral excitatory burst neuron (EBN) area immediately rostral to the abducens nucleus. Systematic tracking for antidromic stimulation revealed a wide distribution of effective spots in and near the EBN area, with varied latencies and thresholds, suggesting terminal branching in that area. The same neurons were also antidromically activated from the contralateral inhibitory burst neuron (IBN) area, the region near the midline, and the nucleus reticularis tegmenti pontis. 3. These neurons exhibited a characteristic firing pattern related to nystagmus: with contralateral rotation the firing rate gradually increased during the slow phase (type II response) and further steeply increased in a burst fashion before and during the contraversive quick phase. Since the time of occurrence of burst activity in these neurons was similar to that of contralateral ENBs and IBNs that received their axonal projection, it is suggested that they send excitatory input to burst neurons, and can thus be called burster-driving neurons (BDNs). 4. Intracellular study revealed that stimulation of the BDN area produced monosynaptic EPSPs in contralateral EBNs. The monosynaptic connection of BDNs with EBNs was confirmed by detecting unitary extracellular synaptic currents of EBNs with the spike-triggered averaging technique. 5. In contrast to BDNs, another class of nystagmus-related type II neurons in the prepositus hypoglossi and medullary reticular formation showed a discharge pattern similar to that of abducens motoneurons on the same side. None of them was antidromically activated from the contralateral pontine reticular formation including the EBN area. Some neurons responded anti-dromically to stimulation of the ipsilateral dorsomedial pontine reticular formation. 6. In conclusion, the input from the horizontal canal during rotation reaches the contralateral prepositus hypoglossi nucleus and the underlying reticular formation through the vestibular nuclei, and a class of neurons in these structures (BDNs) responds to the canal input in a burst fashion following a tonic type II activity. The axons of BDNs cross the midline and monosynaptically excite EBNs on the side of the canal stimulated. The burst activity of BDNs at the quick phase is suggested to contribute to generation of spike burst of EBNs and IBNs.