Hans Straka died in the morning of December 11, 2022 at his home in Munich, unexpected and much too early. He was a dedicated biologist, loved the mountains and was connected to home (Oberammergau, active participant in the Passion Play). His scientific journey took him from Munich via Paris and New York back to Munich and his many academic accomplishments ranged from a membership of the Editorial board of the Journal of Neurophysiology and of the Journal of Neuroscience. He was associate editor for Frontiers in Neuro-otology and for the volume "The Senses" he edited the part on Vestibular Function in 2020. In 2009 he became Professor of Systemic Neurosciences at the Department of Biology in Munich. Apart from his many academic accomplishments, however, Hans was a close friend to those of us who were fortunate enough to get to know him better.

Hans Straka died in the morning of December 11, 2022 at his home in Munich, unexpected and much too early. He was a dedicated biologist, loved the mountains and was connected to home (Oberammergau, active participant in the Passion Play). His scientific journey took him from Munich via Paris and New York back to Munich and his many academic accomplishments ranged from a membership of the Editorial board of the Journal of Neurophysiology and of the Journal of Neuroscience. He was associate editor for Frontiers in Neuro-otology and for the volume "The Senses" he edited the part on Vestibular Function in 2020. In 2009 he became Professor of Systemic Neurosciences at the Department of Biology in Munich. Apart from his many academic accomplishments, however, Hans was a close friend to those of us who were fortunate enough to get to know him better.

Locomotion is associated with a number of optical consequences that degrade visual information processing in the absence of appropriate compensatory movements. The resulting retinal image flow is counteracted by coordinated eye-head reflexes that are initiated by optokinetic and vestibular inputs. The contribution of the vestibulo-ocular reflex (VOR) for stabilizing retinal images is relatively small in amplitude in frogs but important in function by compensating for the non-linearities of the neck motor system. The spatial tuning of the VOR networks underlying the angular (AVOR) and linear (LVOR) with respect to canal and extraocular motor coordinates is organized in a common, canal-related reference frame. Thereby, the axes of head and eye rotation are aligned, principle and auxiliary VOR connections transform vestibular into motor signals and parallel AVOR and LVOR circuits mediate vergence and version signals separately. Comparison of these results with data from other vertebrates demonstrates a number of fundamental organization principles common to most vertebrates. However, the fewer degrees of behavioral freedom of frogs are reflected by the absence of, e.g. a functioning velocity storage network or of a fixation suppression of the VOR. In vitro experiments with the isolated brainstem and branches of N.VIII attached were used to study the putative transmitters of vestibular nerve afferent inputs, the postsynaptic receptor subtypes of second-order vestibular neurons and their dynamic response properties. Evidence is presented that suggests that afferent vestibular nerve fibers with different dynamic response properties activate different subtypes of glutamate receptors. The convergence pattern of monosynaptic afferent nerve inputs from different labyrinthine organs onto second-order vestibular neurons is remarkably specific. As a rule, second-order vestibular neurons receive converging afferent nerve inputs from one semicircular canal and from a specific sector of hair cells on one otolith organ. This convergence pattern remains malleable even in adulthood and reorganization is initiated by activity-related changes in vestibular nerve afferent fibers. The output of second-order vestibular neurons is modified by at least three inhibitory control loops. Uncrossed inhibitory vestibular side loops appear to control specifically the dynamic response tuning, whereas coplanar commissural inhibitory inputs improve mainly the spatial tuning and the cerebellar feedback loop controls the response gain. Among the targets of second-order vestibular projection neurons are extraocular motoneurons and internuclear neurons. Extraocular motoneurons differ among each other by the presence of very different response dynamics. These differences may represent a co-adaptation to the response dynamics of twitch and non-twitch extraocular muscle fibers. Different dynamical properties are required for a rapid acceleration of the globe at the one end and for the maintenance of a stable eccentric eye position over long periods of time at the other end of a continuum of variations in dynamic response properties. The maintenance of a given eccentric eye position over long periods of time is especially well developed in frogs and assists visual surveillance during lurking in the absence of saccades.

Direct and parametric effects of stretch receptors in the extraocular muscles on abducens nerve activity were investigated in the unanesthetized immobilized frog. Horizontal passive rotations of one eye in the physiological range (±5°) did not elicit responses in the activity of abducens nerve on either side; however, larger rotations or pull of one eye evoked long latency direction-unspecific responses simultaneously in both nerves. When the animal was stimulated vestibulary in the horizontal plane with sinusoidal or constant acceleration, abducens activity was not altered in correlation to passive eye movements in the physiological range. Similarly, the activity of either nerve evoked by simultaneous or preceding optokinetic stimulation of one eye with constant pattern velocity was not modified by passive rotation of the contralateral eye.

The ramus anterior (RA) of N.VIII was sectioned unilaterally. Two months later we analyzed in vivo responses of the ipsi- and of the contralesional abducens nerve during horizontal and vertical linear acceleration in darkness. The contralesional abducens nerve had become responsive again to linear acceleration either because of a synaptic reorganization in the vestibular nuclei on the operated side and/or because of a reinnervation of the utricular macula by regenerating afferent nerve fibers. Significant differences in the onset latencies and in the acceleration sensitivities allowed a separation of RA frogs in a group without and in a group with functional utricular reinnervation. Most important, the vector orientation for maximal abducens nerve responses was clearly altered: postlesional synaptic reorganization resulted in the emergence of abducens nerve responses to vertical linear acceleration, a response component that was barely detectable in RA frogs with utricular reinnervation and that was absent in controls. The ipsilesional abducens nerve, however, exhibited unaltered responses in either group of RA frogs. The altered spatial tuning properties of contralesional abducens nerve responses are a direct consequence of the postlesional expansion of signals from intact afferent nerve and excitatory commissural fibers onto disfacilitated 2nd-order vestibular neurons on the operated side. These results corroborate the notion that postlesional vestibular reorganization activates a basic neural reaction pattern with more beneficial results at the cellular than at the network level. However, given that the underlying mechanism is activity-related, rehabilitative training after vestibular nerve lesion can be expected to shape the ongoing reorganization.

Annals of the New York Academy of SciencesVolume 1004, Issue 1 p. 429-433 Article Spatial Convergence Pattern of Canal and Macular Nerve Afferent Signals in Frog Second-Order Vestibular Neurons HANS STRAKA, Corresponding Author HANS STRAKA Physiologisches Institut, LMU München, Pettenkoferstrasse 12, 80336 München, GermanyAddress for correspondence: Dr. H. Straka, Physiologisches Institut, Pettenkoferstrasse 12, 80336 München, Germany. Voice: 49-89-5996-232; fax: 49-89-5996-216. straka@wifomail.med.uni-muenchen.deSearch for more papers by this authorNORBERT DIERINGER, NORBERT DIERINGER Physiologisches Institut, LMU München, Pettenkoferstrasse 12, 80336 München, GermanySearch for more papers by this author HANS STRAKA, Corresponding Author HANS STRAKA Physiologisches Institut, LMU München, Pettenkoferstrasse 12, 80336 München, GermanyAddress for correspondence: Dr. H. Straka, Physiologisches Institut, Pettenkoferstrasse 12, 80336 München, Germany. Voice: 49-89-5996-232; fax: 49-89-5996-216. straka@wifomail.med.uni-muenchen.deSearch for more papers by this authorNORBERT DIERINGER, NORBERT DIERINGER Physiologisches Institut, LMU München, Pettenkoferstrasse 12, 80336 München, GermanySearch for more papers by this author First published: 24 January 2006 https://doi.org/10.1111/j.1749-6632.2003.tb00254.xAboutPDF ToolsRequest permissionExport citationAdd to favoritesTrack citation ShareShare Give accessShare full text accessShare full-text accessPlease review our Terms and Conditions of Use and check box below to share full-text version of article.I have read and accept the Wiley Online Library Terms and Conditions of UseShareable LinkUse the link below to share a full-text version of this article with your friends and colleagues. Learn more.Copy URL Share a linkShare onFacebookTwitterLinkedInRedditWechat Volume1004, Issue1THE OCULOMOTOR AND VESTIBULAR SYSTEMS: THEIR FUNCTION AND DISORDERSOctober 2003Pages 429-433 RelatedInformation

We compared the spatial organization patterns of linear and angular vestibuloocular reflexes in frogs by recording the multiunit spike activity from cranial nerve branches innervating the lateral rectus, the inferior rectus, or the inferior obliquus eye muscles. Responses were evoked by linear horizontal and/or vertical accelerations on a sled or by angular accelerations about an earth-vertical axis on a turntable. Before each sinusoidal oscillation test in darkness, the static head position was systematically altered to determine those directions of horizontal linear acceleration and those planes of angular head oscillation that were associated with minimal response amplitudes. Inhibitory response components during angular accelerations were clearly present, whereas inhibitory response components during linear accelerations were absent. Likewise was no contribution from the vertical otolith organs (i.e., lagena and saccule) observed during vertical linear acceleration. Horizontal linear acceleration evoked responses that originated from eye muscle-specific sectors on the contralateral utricular macula. The sectors of the inferior obliquus and lateral rectus muscles on the utricle had an opening angle of 45 and 60 degrees, respectively and overlapped to a large extent in the laterorostral part of the utricle. Both sectors were coplanar with the horizontal semicircular canals. The sector of the inferior rectus muscle was narrow (opening 5 degrees), laterocaudally oriented, and slightly pitched up by 6 degrees. Angular acceleration evoked maximal responses in the inferior obliquus muscle nerve that originated from the ipsilateral horizontal and the contralateral anterior vertical canals in a ratio of 50:50. Lateral rectus excitation originated from the contralateral horizontal and anterior vertical semicircular canals in a ratio of 80:20. The excitatory responses of the inferior rectus muscle nerve originated exclusively from the contralateral posterior vertical canal. Measured data and known semicircular canal plane vectors were used to calculate the spatial orientation of maximum sensitivity vectors for the investigated eye muscle nerves in semicircular canal coordinates. Comparison of the directions of maximal sensitivity vectors of responses evoked by linear or angular accelerations in a given eye muscle nerve showed that the two vector directions were oriented about orthogonally with respect to each other. With this arrangement the linear and the angular vestibuloocular reflex can support each other dynamically whenever they are co-activated without a change in the spatial response characteristics. The mutual adaptation of angular and linear vestibuloocular reflexes as well as the differences in their organization described here for frogs may represent a basic feature common for vertebrates in general.

Vestibular nerve branches innervating the sensory epithelia of the three semicircular canals or of the three otolith organs of frogs were selectively labeled in-vitro with biocytin. Labeled afferent fibers from the semicircular canals, utricle, and lagena were encountered in each of the four vestibular nuclei and their projections overlapped considerably. Saccular afferent fibers projected to the dorsal (acoustic) nuclei and smaller projections to the vestibular nuclei were regionally restricted. Per semicircular canal or otolith organ about equal numbers (11–14) of medium sized vestibular neurons (between 7.5 and 17 μm in diameter) were dye-coupled to afferent fibers. Most of these dye-coupled vestibular neurons were located in the lateral and descending vestibular nuclei between the VIIIth and IXth nerves. The superior vestibular nucleus was relatively free of dye-coupled vestibular neurons. The location of this subpopulation of central vestibular neurons supports the notion that these neurons are part of a particular vestibulospinal pathway. In addition, from each of the canal and/or otolith organs about 3–4 efferent vestibular neurons were labeled retrogradely. These neurons (between 15 and 26 μm in diameter) were located ventral to the vestibular nuclear complex. The branching of efferent vestibular neurons was shown by the presence of neurons that were double labeled by two different fluorescent dyes applied in the same experiment to the anterior and posterior ramus of the same VIIIth nerve, respectively. The branching of these efferent neuron axons explained the presence of collaterals and terminals in the sensory epithelia of a number of untreated ipsilateral endorgans.

Nerve injury induces a reorganization of subcortical and cortical sensory or motor maps in mammals. A similar process, vestibular plasticity 2 mo after unilateral section of the ramus anterior of N. VIII was examined in this study in adult frogs. The brain was isolated with the branches of both N. VIII attached. Monosynaptic afferent responses were recorded in the vestibular nuclei on the operated side following ipsilateral electric stimulation either of the sectioned ramus anterior of N. VIII or of the intact posterior vertical canal nerve. Excitatory and inhibitory commissural responses were evoked by separate stimulation of each of the contralateral canal nerves in second-order vestibular neurons. The afferent and commissural responses of posterior vertical canal neurons recorded on the operated side were not altered. However, posterior canal-related afferent inputs had expanded onto part of the deprived ramus anterior neurons. Inhibitory commissural responses evoked from canal nerves on the intact side were detected in significantly fewer deprived ramus anterior neurons than in controls, but excitatory commissural inputs from the three contralateral canal nerves had expanded. This reactivation might facilitate the survival of deprived neurons and reduce the asymmetry in bilateral resting activities but implies a deterioration of the original spatial response tuning. Extensive similarities at the synaptic and network level were noted between this vestibular reorganization and the postlesional cortical and subcortical reorganization of sensory representations in mammals. We therefore suggest that nerve injury activates a fundamental neural reaction pattern that is common between sensory modalities and vertebrate species.

The anterior branch of N. VIII was sectioned in adult frogs. Two months later the brain was isolated to record in vitro responses in the vestibular nuclei and from the abducens nerves following electric stimulation of the anterior branch of N. VIII or of the posterior canal nerve. Extra- and intracellularly recorded responses from the intact and operated side were compared with responses from controls. Major changes were detected on the operated side: the amplitudes of posterior canal nerve evoked field potentials were enlarged, the number of vestibular neurons with a monosynaptic input from the posterior canal nerve had increased, and posterior canal nerve stimulation recruited stronger abducens nerve responses on the intact side than vice versa. Changes in the convergence pattern of vestibular nerve afferent inputs on the operated side strongly suggest the expansion of posterior canal-related afferent inputs onto part of those vestibular neurons that were deprived of their afferent vestibular input. As a mechanism we suggest reactive synaptogenesis between intact posterior canal afferent fibers and vestibularly deprived second-order vestibular neurons.

Most second-order vestibular neurons receive a canal-specific monosynaptic excitation, although the central projections of semicircular canal afferents overlap extensively. This remarkable canal specificity prompted us to study the spatial organization of evoked field potentials following selective stimulation of individual canal nerves. Electrically evoked responses in the vestibular nuclei were mapped systematically in vitro. Constructed activation maps were superimposed on a cytoarchitectonically defined anatomical map. The spatial activation maps For pre- and postsynaptic response components evoked by stimulation of a given canal nerve were similar. Activation maps for monosynaptic inputs from different canals tended to show a differential distribution of their peak amplitudes, although the overlap was considerable. Anterior vertical canal signals peaked in the superior vestibular nucleus, posterior vertical canal signals peaked in the descending and in the dorsal part of the lateral vestibular nucleus, whereas horizontal canal signals peaked in the descending and in the ventral part of the lateral vestibular nucleus. A similar, differential but overlapping, spatial organization of the canal inputs was described also for other vertebrates. suggesting a crude but rather conservative topographical organization of semicircular canal nerve projections within the vestibular nuclei. Differences in the precision of topological representations between vestibular and other sensory modalities are discussed. (C) 2000 Elsevier Science B.V. All rights reserved.

Removal of the labyrinthine organs on one side results in a number of severe postural and dynamic reflex deficits. Over time some of these behavioral deficits normalize again. At a chronic stage the brain of frogs exhibits a number of changes in vestibular and propriospinal circuits on the operated side that were studied in vitro. The onset of changes in the vestibular nuclear complex was delayed, became evident only after head posture had recovered by more than 50%, and was independent of the presence or absence of a degeneration of vestibular nerve afferent fibers. The time course of changes measured in the isolated spinal cord paralleled the time course of normalization of head and body posture. Results obtained after selective lesions of individual labyrinthine nerve branches show that unilateral inactivation of utricular afferent inputs is a necessary and sufficient condition to provoke postural deficits and propriospinal changes similar to those after the removal of all labyrinthine organs. The presence of multiple synaptic changes at distributed anatomic sites over different periods of time suggests that different parts of the central nervous system are involved in the normalization of different manifestations of the vestibular lesion syndrome.

Static and dynamic otolith, horizontal vestibular and optokinetic ocular reflexes were investigated in pigmented rats 1–6 and more months after unilateral vestibular nerve (UVN) section. Evoked responses were compared with published data from control rats studied under identical conditions. Static lateral tilt of UVN rats in the light evoked a vertical deviation in static eye position that was as large as in controls. In darkness, the evoked responses in UVN rats 6 months after the lesion were consistently smaller than in controls. Linear horizontal acceleration in darkness evoked vertical and torsional response components in UVN rats that were parallel-shifted towards lower gains and larger phase lags. Off-vertical axis rotation on a platform provoked responses that differed markedly from those recorded in intact rats with respect to the bias velocity component. These results suggest a permanent deficiency in the static and dynamic otolith-ocular reflex performance of UVN rats. Ocular responses to horizontal table velocity steps in darkness exhibited a direction-specific asymmetry in UVN rats. Step responses evoked by acceleration towards the intact side were larger in gain and longer in duration than responses evoked by acceleration towards the operated side. When compared with control data, responses to either side were reduced in UVN rats and the velocity store mechanism was barely activated by velocity steps towards the operated side. Responses evoked by horizontal optokinetic stimulation with constant pattern velocities were below control values in either direction. Slow-phase eye velocity saturated at much lower values than in intact rats, particularly during pattern motion towards the intact side. The duration of the optokinetic afternystagmus was asymmetrically reduced with respect to control data. Practically identical reductions in duration were found for vestibulo-ocular responses in the opposite directions. Behaving animals exhibited no obvious impairment in their spontaneous locomotory or exploratory activities. However, each UVN rat was impaired, even 2 years after the lesion, in its postural reaction to being lifted by the tail in the air. This observation suggests the presence of a permanent deficit in static and dynamic otolith-spinal reflexes that may be substituted on the ground by proprioceptive inputs.

Second-order vestibular neurons (secondary VNs) were identified in the in vitro frog brain by their monosynaptic excitation following electrical stimulation of the ipsilateral VIIIth nerve. Ipsilateral disynaptic inhibitory postsynaptic potentials were revealed by bath application of the glycine antagonist strychnine or of the gamma-aminobutyric acid-A (GABA(A)) antagonist bicuculline. Ipsilateral disynaptic excitatory postsynaptic potentials (EPSPs) were analyzed as well. The functional organization of convergent monosynaptic and disynaptic excitatory and inhibitory inputs onto secondary VNs was studied by separate electrical stimulation of individual semicircular canal nerves on the ipsilateral side. Most secondary VNs (88%) received a monosynaptic EPSP exclusively from one of the three semicircular canal nerves; fewer secondary VNs (10%) were monosynaptically excited from two semicircular canal nerves; and even fewer secondary VNs (2%) were monosynaptically excited from each of the three semicircular canal nerves. Disynaptic EPSPs were present in the majority of secondary VNs (68%) and originated from the same (homonymous) semicircular canal nerve that activated a monosynaptic EPSP in a given neuron (22%), from one or both of the other two (heteronymous) canal nerves (18%), or from all three canal nerves (28%). Homonymous activation of disynaptic EPSPs prevailed (74%) among those secondary VNs that exhibited disynaptic EPSPs. Disynaptic inhibitory postsynaptic potentials (IPSPs) were mediated in 90% of the tested secondary VNs by glycine, in 76% by GABA, and in 62% by GABA as well as by glycine. These IPSPs were activated almost exclusively from the same semicircular canal nerve that evoked the monosynaptic EPSP in a given secondary VN. Our results demonstrate a canal-specific, modular organization of vestibular nerve afferent fiber inputs onto secondary VNs that consists of a monosynaptic excitation from one semicircular canal nerve followed by disynaptic excitatory and inhibitory inputs originating from the homonymous canal nerve. Excitatory and inhibitory second-order (secondary) vestibular interneurons are envisaged to form side loops that mediate spatially similar but dynamically different signals to secondary vestibular projection neurons. These feedforward side loops are suited to adjust the dynamic response properties of secondary vestibular projection neurons by facilitating or disfacilitating phasic and tonic input components.

This study describes the localization of γ-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 a 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 or glycine immunoreactive cells revealed a complementatory 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 vestibulo-ocular 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. J. Comp. Neurol. 377:149–164, 1997. © 1997 Wiley-Liss, Inc.

The neuronal organization of the horizontal vestibulo-ocular reflex (VOR) of frogs is reviewed and compared to results obtained from other vertebrates. Extensive similarities in the anatomical, physiological and pharmacological properties of the basic, three-neuronal reflex arc indicate the presence of a conservative, common plan for the VOR of vertebrates and its inhibitory control. To meet the requirements resulting from different locomotor patterns and from misalignments between semicircular canal and extraocular muscle planes, species-specific adaptations of the VOR performance must be present as well. Major differences in the performance of supplementary networks in different species provide a basis for this coadaptation.

1. Eighth nerve evoked responses in central vestibular neurons (n = 146) were studied in the isolated brain stem of frogs. Ninety percent of these neurons responded with a monosynaptic excitatory postsynaptic potential (EPSP) after electrical stimulation of the ipsilateral VIIIth nerve. In 5% of these neurons, the EPSP was truncated by a disynaptic inhibitory postsynaptic potential (IPSP), and in 5% of these neurons a pure disynaptic IPSP was evoked. 2. Disynaptic IPSPs superimposed upon apparently pure EPSPs were revealed by bath application of the glycine receptor antagonist strychnine (0.5–5 microM) or of the gamma-aminobutyric acid-A (GABAA) receptor antagonist bicuculline (0.5–2 microM). The evoked EPSP increased in most central vestibular neurons (strychnine: 15 out of 16 neurons; bicuculline 26 out of 29 neurons). At higher stimulus intensities, the evoked spike discharge increased from 2 to 3 spikes before up to 8-10 spikes per electrical pulse during the application of blocking agents. The unmasked disynaptic inhibitory component increased with stimulus intensity to a different extent in different neurons. 3. Lesion studies demonstrated that these inhibitory components were generated ipsilaterally with respect to the recording side. The disynaptic strychnine-sensitive inhibition was mediated by neurons located either in the ventral vestibular nuclear complex (VNC) or in the adjacent reticular formation. The spatial distribution of the disynaptic inhibition was investigated by simultaneous recordings of VIIIth nerve-evoked field potentials at different rostrocaudal locations of the VNC. A significant strychnine-sensitive component was detected in the middle and caudal parts but not in the rostral part of the VNC. A bicuculline-sensitive component was detected in the rostral and in the caudal parts but not in the middle part of the VNC. In view of a similar rostrocaudal distribution of glycineor GABA-immunoreactive neurons in the VNC of frogs, our results suggest that part of the disynaptic inhibition is mediated by local interneurons with a spatially restricted projection area. 4. The monosynaptic EPSP of second-order vestibular neurons was mediated in part by N-methyl-D-aspartate (NMDA) and in part by non-NMDA receptors. The relative contribution of the NMDA receptor-mediated component of the EPSP decreased with stronger stimuli. This negative correlation could have resulted from a preferential activation of NMDA receptors via thick vestibular nerve afferent fibers. Alternatively, the activation of NMDA receptors became disfacilitated at higher stimulus intensities due to the recruitment of disynaptic inhibitory inputs. Comparison of data obtained in the presence and in the absence of these glycine and GABAA receptor blockers indicates a preferential activation of NMDA receptors via larger-diameter vestibular nerve afferent fibers. 5. The kinetics of NMDA receptors (delay, rise time) activated by afferent nerve inputs were relatively fast. These fast kinetics were independent of superimposed IPSPs. The association of these receptors with large-diameter vestibular nerve afferent fibers suggests that fast NMDA receptor kinetics might be matched to the more phasic response dynamics of the large diameter vestibular afferent neurons to natural head accelerations.