Saccadic omnipause neurons (OPNs) are essential for the generation of saccadic eye movements. In primates OPNs are located near the midline within the nucleus raphe interpositus (rip). In the present study we used several different neuroanatomical methods to investigate the transmitters associated with OPNs in the monkey. Immunolabeling for the calcium-binding protein parvalbumin was employed to mark OPNs in the monkey and define the homologous cell group in cat and human. The use of antibodies against GABA, glycine (GLY), glutamate (GLU), serotonin (5-HT), and tyrosine hydroxylase revealed that the somata of OPNs are GLY immunoreactive, but they are devoid of GABA and 5-HT immunostaining. In situ hybridization with the GAD67 mRNA probe confirmed the negative GABA immunostaining of OPNs. 3H-GLY was injected into a projection field of OPNs, the rostral interstitial nucleus of the medial longitudinal fascicle (riMLF)--the vertical saccadic burst neuron area. This resulted in selective retrograde labeling of the OPNs in rip, while no labeling was found in the superior colliculus, which sends an excitatory projection to the riMLF. The somata and dendrites of putative burst neurons in the riMLF were contacted by numerous GLY- immunoreactive terminals. The quantitative analysis of immunoreactive terminal-like structures contacting OPNs revealed a strong input from GLY- and GABA-positive terminals on somata and dendrites, whereas GLU- positive puncta were mainly confined to the dendrites. Very few 5-HT and catecholaminergic terminals contacted OPN somata. Our findings suggest that OPNs use GLY as a neurotransmitter, and they receive numerous contacts from GABAergic, glycinergic, and glutaminergic afferents, and significantly fewer from monoaminergic inputs.

Central vestibular neurons receive substantial inputs from the contralateral labyrinth through inhibitory and excitatory brainstem commissural pathways. The functional organization of these pathways was studied by a multi-methodological approach in isolated frog whole brains. Retrogradely labeled vestibular commissural neurons were primarily located in the superior vestibular nucleus in rhombomeres 2/3 and the medial and descending vestibular nucleus in rhombomeres 5-7. Restricted projections to contralateral vestibular areas, without collaterals to other classical vestibular targets, indicate that vestibular commissural neurons form a feedforward pushpull circuitry. Electrical stimulation of the contralateral coplanar semicircular canal nerve evoked in canal-related second-order vestibular neurons (2°VN) commissural IPSPs (ϳ70%) and EPSPs (ϳ30%) with mainly (ϳ70%) disynaptic onset latencies. The dynamics of commissural responses to electrical pulse trains suggests mediation predominantly by tonic vestibular neurons that activate in all tonic 2°VN large-amplitude IPSPs with a reversal potential of Ϫ74 mV. In contrast, phasic 2°VN exhibited either nonreversible, smallamplitude IPSPs (ϳ40%) of likely dendritic origin or large-amplitude commissural EPSPs (ϳ60%). IPSPs with disynaptic onset latencies were exclusively GABAergic (mainly GABA A receptor-mediated) but not glycinergic, compatible with the presence of GABAimmunopositive (ϳ20%) and the absence of glycine-immunopositive vestibular commissural neurons. In contrast, IPSPs with longer, oligosynaptic onset latencies were GABAergic and glycinergic, indicating that both pharmacological types of local inhibitory neurons were activated by excitatory commissural fibers. Conservation of major morpho-physiological and pharmacological features of the vestibular commissural pathway suggests that this phylogenetically old circuitry plays an essential role for the processing of bilateral angular head acceleration signals in vertebrates.

Labyrinthine nerve-evoked monosynaptic excitatory postsynaptic potentials (EPSPs) in second-order vestibular neurons (2 degrees VN) sum with disynaptic inhibitory postsynaptic potentials (IPSPs) that originate from the thickest afferent fibers of the same nerve branch and are mediated by neurons in the ipsilateral vestibular nucleus. Pharmacological properties of the inhibition and the interaction with the afferent excitation were studied by recording monosynaptic responses of phasic and tonic 2 degrees VN in an isolated frog brain after electrical stimulation of individual semicircular canal nerves. Specific transmitter antagonists revealed glycine and GABA(A) receptor-mediated IPSPs with a disynaptic onset only in phasic but not in tonic 2 degrees VN. Compared with GABAergic IPSPs, glycinergic responses in phasic 2 degrees VN have larger amplitudes and a longer duration and reduce early and late components of the afferent nerve-evoked subthreshold activation and spike discharge. The difference in profile of the disynaptic glycinergic and GABAergic inhibition is compatible with the larger number of glycinergic as opposed to GABAergic terminal-like structures on 2 degrees VN. The increase in monosynaptic excitation after a block of the disynaptic inhibition in phasic 2 degrees VN is in part mediated by a N-methyl-d-aspartate receptor-activated component. Although inhibitory inputs were superimposed on monosynaptic EPSPs in tonic 2 degrees VN as well, the much longer latency of these IPSPs excludes a control by short-latency inhibitory feed-forward side-loops as observed in phasic 2 degrees VN. The differential synaptic organization of the inhibitory control of labyrinthine afferent signals in phasic and tonic 2 degrees VN is consistent with the different intrinsic signal processing modes of the two neuronal types and suggests a co-adaptation of intrinsic membrane properties and emerging network properties.

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Horizontal eye movements in response to vestibular and optokinetic stimulation were investigated in 20 species of marine and fresh water teleosts. Most species performed spontaneous saccadic eye movements in light and in darkness. Successive saccades occurred either at irregular intervals and in different directions (few species), or in bursts of smaller saccades in one direction followed by saccades in the opposite direction (majority of species). Thus, in the latter group the lines of sight shifted cyclically from one side to the other about every 10–15 s. Eye movement responses in goldfish and toadfish were prototypical for two major functional groups. Optokinetic and vestibular nystagmus in the goldfish were much more regular and attained significantly higher gains than in the oyster toadfish. Optokinetic afternystagmus (OKAN) and perVpostrotatory nystagmus lasted about 10–12 s in the goldfish, but only 2–3 s in the oyster toadfish. Bilateral labyrinthectomy in goldfish resulted in optokinetic gains and OKAN durations that were very close to those of intact toadfish. These results indicate the presence (in goldfish) and the absence (in toadfish) of a functioning velocity storage network, respectively. Other non-ostariophysan teleosts failed to show a ''goldfish-like'' response pattern as well. Ostariophysan species differed in their reflex performance as well. A ''gold-fish-like'' response pattern was observed in five of the seven cypriniform species and in two of the four characiform species but not in the four siluriform species. Positively correlated, combined response properties suggest that some species are better equipped for gaze stabilization at higher velocities than other species. These behavioral differences may parallel different specializations in locomotor pattern and mode of life. Absence of a functioning velocity storage network in bottom-dwelling teleosts (as in Amphibia) may be related to the sporadic, slow locomotion of these species and the resulting small requirements for continuous gaze stabilization during self-motion at higher velocities.