Topographic representation of vestibular and somatosensory signals in the anuran thalamus.

In the isolated brain of the fire-bellied toad, Bombina orientalis, the spatial distribution of vestibular and somatosensory responses in thalamic nuclei was studied following electrical activation of the Vth nerve, the ramus anterior of the VIIIth nerve and of the dorsal roots of spinal nerves 3 and 8. Responses were systematically mapped in frontal planes through the diencephalon at four rostro-caudal levels. The calculated activity maps were superimposed on the outlines of diencephalic nuclei, and those nuclei that received particularly large inputs from the stimulated sensory nerve roots were indicated. Maximal response amplitudes coincided with ventral, central, and posterior thalamic areas and exhibited a topography that differed for each sensory nerve root. Maximal responses evoked from the Vth nerve were largely separated from those from spinal dorsal roots 3 and 8, whereas maximal vestibular responses partly overlapped with those from the other somatosensory nerve roots. Our findings indicate that within the amphibian thalamus sensory signals originating from different nerve roots are largely represented in separate areas as is the case in the thalamus of amniotes. However, the anterior dorsal thalamus which is the only origin of ascending pathways to the medial and dorsal pallium (assumed homologues of the mammalian hippocampus and neocortex, respectively) receives only minor vestibular and somatosensory input. This corroborates the view that amphibians lack a direct sensory thalamo-cortical, or "lemnothalamic," pathway typical of mammals and birds.

Distribution of GABA, glycine, and glutamate immunoreactivities in the vestibular nuclear complex of the frog.

This study describes the localization of gamma-aminobutyric acid (GABA), glycine, and glutamate immunoreactive neurons, fibers, and terminal-like structures in the vestibular nuclear complex (VNC) of the frog by using a postembedding procedure with consecutive semithin sections at the light microscopic level. For purposes of this study, the VNC was divided into a medial and lateral region. Immunoreactive cells were observed in all parts of the VNC. GABA-positive neurons, generally small in size, were predominantly located in the medial part of the VNC. Glycine-positive cells, more heterogeneous in size than GABA-positive cells, were scattered throughout the VNC. A quantitative analysis of the spatial distribution of GABA glycine immunoreactive cells revealed a complementary relation between the density of GABA and glycine immunoreactive neurons along the rostrocaudal extent of the VNC. In about 10% of the immunolabeled neurons, GABA and glycine were colocalized. Almost all vestibular neurons were, to a variable degree, glutamate immunoreactive, and colocalization of glutamate with GABA and/or glycine was typical. GABA, glycine, or glutamate immunoreactive puncta were found in close contact to somata and main dendrites of vestibular neurons. A quantitative analysis revealed a predominance of glutamate-positive terminal-like structures compared to glycine or GABA containing profiles. A small proportion of terminal-like structures expressed colocalization of GABA and glycine or glycine and glutamate. The results are compared with data from mammals and discussed in relation to vestibuloocular and vestibulo-spinal projection neurons, and vestibular interneurons. GABA and glycine are the major inhibitory transmitters of these neurons in frogs as well as in mammals. The differential distribution of GABA and glycine might reflect a compartmentalization of neurons that is preserved to some extent from the early embryogenetic segmentation of the hindbrain.

Electrophysiological and pharmacological characterization of vestibular inputs to identified frog abducens motoneurons and internuclear neurons in vitro.

Synaptic vestibular inputs of antidromically identified motoneurons and internuclear neurons in the abducens nucleus were studied electrophysiologically and pharmacologically in the isolated brain of grass frogs (Rana temporaria). The prevailing response pattern of abducens motoneurons (AbMOT) following stimulation of the VIIIth nerve was crossed disynaptic excitation and uncrossed disynaptic inhibition. A few AbMOT (five of 46), however, exhibited uncrossed excitation instead of inhibition. Abducens internuclear neurons (AbINT), identified by antidromic activation following stimulation of the contralateral medial longitudinal fascicle, exhibited disynaptic response patterns to stimulation of the VIIIth nerve that were very similar in latency and rise time to those of AbMOT except for the absence of uncrossed disynaptic inhibition. Bath application of strychnine (50 microM), a glycine antagonist, blocked the uncrossed inhibitory vestibular input to AbMOT and AbINT completely and reversibly, whereas picrotoxin (100 microM), a GABA (gamma-aminobutyric acid) antagonist, had no detectable effect on these disynaptic potentials. These results suggest glycine as the transmitter of inhibitory vestibular projections onto AbMOT and AbINT. The pharmacology of the excitatory vestibular input of these neurons was studied by electrical stimulation of the vestibular nuclear complex. Crossed monosynaptic excitatory inputs in AbMOT and AbINT were blocked completely by CNQX (6-cyano-7-nitroquinoxaline-2,3-dione) (10 microM), an antagonist of AMPA (alpha-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid) receptors, indicating glutamatergic excitation. Comparison of these results with those in the cat suggests the presence of a basic horizontal vestibulo-ocular reflex that is very similarly organized, and corroborates the hypothesis that major behavioural differences in the performance of compensatory eye movements between species result from the properties of supplementary networks and not from differences in a common 'three-neuron' vestibulo-ocular arc.