Neuronal activity in the vestibular nuclei of the alert monkey during vestibular and optokinetic stimulation

Summary Recordings from neurons of the vestibular nuclei were performed in alert monkeys. Type I and type II units were identified by rotating the monkey about a vertical axis. All neurons responded also when only the visual surround was rotated around the stationary monkey. The combination of visual and vestibular stimulation points towards non-algebraic summation characteristics for the two inputs, with each input dominating the response over a certain range.Supported by Swiss National Foundation for Scientific Research 3.044.76 and Emil-Barell-Foundation of Hoffmann-La Roche, Basel, Switzerland     

Linear addition of optokinetic and vestibular signals in the vestibular nucleus

Summary A simple model of the vestibuloocular reflex and the optokinetic system was used to simulate recent data on visual and vestibular responses of neurons in the vestibular nucleus. Contrary to a previous interpretation, the results support the hypothesis that the optokinetic and semicircular canal signals are combined simply by linear addition on the cells of the vestibular nucleus.     

Vestibular nuclei activity during optokinetic after-nystagmus (OKAN) in the alert monkey

Summary Neurons which receive an input from the horizontal semicircular canals were recorded from the vestibular nuclei in chronically prepared monkeys (Macaca mulatta) during optokinetic after-nystagmus (OKAN). In complete darkness the vestibular neurons showed activity changes which closely paralleled the strength of nystagmus. The activity of vestibular units returned to baseline levels of spontaneous discharge only when all after-nystagmus had ceased, or when it was inhibited by stationary visual stimuli. The possible role of vestibular neurons in the generation of OKAN and its significance in vestibulo-visual interaction is discussed.Supported in part by Swiss National Foundation for Scientific Research 3.672-0.77 and Emil Barell-Foundation of Hoffman-La Roche, Basel, Switzerland     

Vestibular nuclei activity in the alert monkey during suppression of vestibular and optokinetic nystagmus

Summary Single neurons were recorded in the vestibular nuclei of monkeys trained to suppress nystagmus by visual fixation during vestibular or optokinetic stimulation. During optokinetic nystagmus vestibular nuclei neurons exhibit frequency changes. With the suppression of optokinetic nystagmus this neuronal activity on average is attenuated by 40% at stimulus velocities of 40 °/s. At a stimulus velocity of 5 °/s responses are, under both conditions, close to threshold. For steps in velocity, suppression of vestibular nystagmus shortens the time constants of the decay of neuronal activity from 15–35 s to 5–9 s, while the amplitude of the response remains unchanged. The results are discussed in relation to current models of visual-vestibular interaction. These models use a feedback mechanism which normally operates during vestibular and optokinetic nystagmus. Nystagmus suppression interrupts this feedback loop.Supported by the Swiss National Foundation for Scientific Research (SNF 3.233.77) and the Deutsche Forschungsgemeinschaft (U.W. Buettner, Bue 379/2)     

Thioperamide delays vestibular compensation in goldfish

Unilateral lesion of the vestibular system induces posturo-locomotor deficits that are compensated for with time. Drug therapy is currently used to improve the recovery process and to facilitate vestibular compensation. We investigated the effects of thioperamide on functional recovery after unilateral labyrinthectomy in Carassius auratus. Approximately 24h after surgery, the animals were injected intraperitoneally with thioperamide (15 mg/kg) and saline (1.5 ml/kg). The injections were repeated daily for a total of 15 consecutive days. The substances were administered in a volume of 1.5 ml/kg body weight. Another group, which served as a non-lesion control, did not receive unilateral labyrinthectomy or system injections. Animals treated with saline presented a compensatory decrease in body tilt on the 7th day, while the animals treated with thioperamide presented a decrease in body tilt from the 13th day, suggesting a delay in the functional recovery process. These results suggest that an increase in cerebral histamine levels impairs vestibular compensation in goldfish.     

Persistence of visual response in vestibular nucleus neurons in cerebellectomized cat

Summary Responses to pure visual stimuli and visual-vestibular interactions similar to those reported in other vertebrates were found in single vestibular nucleus neurons in cat. Contrary to prevailing hypotheses, the cerebellum is not an essential part of the pathway mediating these visual responses.E.L. Keller was partially supported by a grant from the Alexander von Humboldt Foundation and NIH grant EY 0095S-07 while on leave from the Dept. of Electrical Engineering and Computer Sciences, Univ. of California, Berkeley     

Nystagmus induced by electrical stimulation of the vestibular and prepositus hypoglossi nuclei in the monkey: evidence for site of induction of velocity storage

Summary Electrical stimulation of the vestibular nuclei (VN) and prepositus hypoglossi nuclei (PPH) of alert cynomolgus monkeys evoked nystagmus and eye deviation while they were in darkness. At some sites in VN, nystagmus and after-nystagmus were induced with characteristics suggesting that velocity storage had been excited. We analyzed these responses and compared them to the slow component of optokinetic nystagmus (OKN) and to optokinetic after-nystagmus (OKAN). We then recorded unit activity in VN and determined which types of nystagmus would be evoked from the sites of recording. Nystagmus and eye deviations were also elicited by electrical stimulation of PPH, and we characterized the responses where unit activity was recorded in PPH. Horizontal slow phase velocity of the VN storage responses was contralateral to the side of stimulation. The rising time constants and peak steady-state velocities were similar to those of OKN, and the falling time constants of the after-nystagmus and of OKAN were approximately equal. Both the induced after-nystagmus and OKAN were habituated by stimulation of the VN. When horizontal after-nystagmus was evoked with animals on their sides, it developed yaw and pitch components that tended to shift the vector of the slow phase velocity toward the spatial vertical. Similar cross-coupling occurs for horizontal OKAN or for vestibular post-rotatory nystagmus elicited in tilted positions. Thus, the storage component of nystagmus induced by VN stimulation had the same characteristics as the slow component of OKN and the VOR. Positive stimulus sites for inducing nystagmus with typical storage components were located in rostral portions of VN. They lay in caudal ventral superior vestibular nucleus (SVN), dorsal portions of central medial vestibular nucleus (MVN) caudal to the abducens nuclei and in adjacent lateral vestibular nucleus (LVN). More complex stimulus responses, but with contralateral after-nystagmus, were induced from surrounding regions of ventral MVN and LVN, rostral descending vestibular nucleus and the marginal zone between MVN and PPH. Vestibular-only (VO), vestibular plus saccade (VPS) and tonic vestibular pause (TVP) units were identified by extracellular recording. Stimulation near type I lateral and vertical canalrelated VO units elicited typical storage responses with after-nystagmus in 23 of 29 tracks (79%). Stimulus responses were more complex from the region of neurons with oculomotor-related signals, i.e., TVP or VPS cells, although after-nystagmus was also elicited from these sites. Effects of vestibular nerve and nucleus stimulation were compared. Nerve stimulation evoked nystagmus with both a rapid and slow component and after-nystagmus. There was a more prominent rapid rise in slow phase velocity, higher peak velocities, shorter latencies and a shorter falling time constant from nerve than from nucleus stimulation. This indicates more prominent activation of rapid pathways from nerve stimulation. From a comparison of nerve- and nucleus-induced nystagmus, we infer that there was predominant activation of the network responsible for velocity storage by electrical stimulation at many sites in the VN. Microstimulation at sites in PPH elicited nystagmus with ipsilateral slow phases or ipsilateral eye deviations. Slow phase eye velocity changed rapidly at the onset of nystagmus, and peak eye velocities were about 10–15°/s lower than from VN stimulation. The nystagmus had no slow component, and it was not followed by after-nystagmus. Only burst or burst-tonic neurons were recorded in PPH. Stimulation at sites of recording of these units induced either nystagmus with a rapid component or ipsilateral eye deviation. We conclude that the slow component of optokinetic and vestibular nystagmus, attributable to velocity storage is produced in the VN, not in the PPH. We postulate that VO neurons lying in caudal ventral portions of SVN, dorsal portions of MVN and adjacent LVN are part of the network that generates velocity storage.     

The vestibular nuclei in the domestic hen

Summary Stereotaxic injections of small quantities of horseradish peroxidase (HRP) are made in the vestibular complex in the hen and the labelled cells due to retrograde transport of the tracer enzyme are studied, especially in the vestibular complex but also in other parts of the brain stem. The superior vestibular nucleus sends commissural fibres to the superior, medial, and descending nuclei and to Deiters' complex. The cell group A projects to the contralateral superior nucleus and to the Deiters' complex. The medial nucleus projects to the contralateral superior and descending nuclei, Deiters' complex, as well as strongly to the medial nucleus. The descending nucleus projects commissurally to the superior nucleus, the medial nucleus and the Deiters' complex, as well as heavily to the descending nucleus. The nucleus Deiters ventralis, the nucleus Deiters dorsalis, the cell group B and the tangential nucleus do not project to any other vestibular nuclei. Furthermore, the medial nucleus projects to the superior and descending nuclei and the Deiters' complex on the same side. The descending nucleus projects to the superior and medial nuclei on the same side. Finally, the superior nucleus, the medial nucleus, the descending nucleus, and the Deiters' complex receive fibres from the ipsilateral nucleus of Darkschewitsch and the nucleus ectomammillaris, as well as bilaterally from the pontine but mainly from the bulbar reticular formation.     

Properties of utricular-activated vestibular neurons that project to the contralateral vestibular nuclei in the cat

The properties of utricular (UT)-activated vestibular neurons that send axons to the contralateral vestibular nuclei (commissural neurons) were investigated intracellularly or extracellularly in decerebrate cats. A total of 27 vestibular neurons were orthodromically activated by stimulation of UT nerves and antidromically activated by stimulation of the contralateral vestibular nuclei. All neurons tested were classified as vestibulospinal (VS), vestibulooculospinal (VOS), vestibuloocular (VO), and unidentified vestibular neurons (V) after antidromic stimulation of the spinal cord and oculomotor/trochlear nuclei. Most UT-activated commissural neurons (20/27) received monosynaptic inputs. Twelve of 27 commissural neurons were located in the medial vestibular nucleus, 5 were in the lateral vestibular nucleus, 10 were in the descending vestibular nucleus, and no commissural neurons were recorded in the superior vestibular nucleus. Seven of 27 neurons were commissural VS neurons, 9 of 27 were commissural VOS neurons, and 11 of 27 were commissural V neurons. No commissural VO neurons were found. All VOS neurons and 3 VS neurons issued descending axons via the medial vestibulospinal tract. We also studied convergent inputs from the posterior semicircular canal (PC) nerve onto UT-activated commissural neurons. Five of 27 UT-activated commissural neurons received converging inputs from the PC nerves. Electronic Publication     

Visual modulation of otolith-dependent units in cat vestibular nuclei

Summary Vestibular nucleus units responding to translational self-motion are shown to respond also to translational movement of a large visual field. These visual-vestibular interactions in otolith-dependent units are similar to those found in canal-related units and could provide the basis for linearvection.     

Visual-vestibular interaction studied with stroboscopically illuminated visual patterns

Summary Horizontal DC-electrooculograms were recorded in subjects rotating on a horizontal turntable sinusoidally at 0.1 Hz and 35 to 40° amplitude. The subjects either fixated a stroboscopically illuminated vertically striped pattern (1.15 to 3.45° period) rotating with the turntable or initiated Sigma-OKN before the rotation began and tried to maintain Sigma-OKN during rotation. In a third paradigm, interaction of vestibulo-ocular reflex (VOR) and Phi OKN was studied. (1) VOR-suppression by fixation was complete within the limits of EOG-recording precision (±1° · s^-1) for flash frequencies f_s>10 flashes · s^-1. VOR-suppression decreased monotonically with f_s between 10 and 1 flashes · s^-1. (2) A similar dependency on f_s was found for VOR-suppression during Sigmaor Phi-OKN. Above 10 flashes · s^-1 VOR-suppression remained incomplete; below 5 flashes · s^-1 VOR-suppression was stronger with the Sigma-OKN paradigm than during fixation and depended on spatial frequency of the pattern. (3) During sinewave rotation of the subject the perceived speed V_p of Sigma-movement correlated to the movement of gaze in space and not to the movement of the eye in head. (4) In a control experiment with normal optokinetic stimulation, OKN-suppression by fixating a small flashing target was found to depend on f_s in a similar way as VOR-suppression in the experiments described above.     

The vestibular nuclei in the domestic hen (Gallus domesticus). I. Normal anatomy.

Summary The topography and the cyto-and fiber architecture of the vestibular nuclear complex in the domestic hen are described as seen in transverse and horizontal thionine and myelin impregnated sections. The subdivision of the nuclear complex arrived at from these studies is discussed in the light of some experimental studies of the fiber connections of the vestibular nuclei in birds and compared with the well known organization of the vestibular nuclei in mammals.Six main vestibular nuclei are identified, the superior nucleus, the nucleus Deiters ventralis, the nucleus Deiters dorsalis, the tangential nucleus, the medial nucleus and the descending nucleus. In addition two cell groups (the cell group A and B) lying in close relation to the other nuclei are described and considered as parts of the vestibular complex. The map of the vestibular complex arrived at is largely in agreement with the maps presented by most earlier authors on other species. Furthermore, it appears that the organization of the vestibular complex in birds is more similar to the organization of the complex in mammals than hitherto recognized.     

Vestibular and somatosensory interaction in the cat vestibular nuclei

Summary The vestibular nuclei of cats were explored extracellularly with micropipettes to locate units with a resting discharge rate which responded to rotation in the horizontal plane. These units were examined for somatosensory input from neck and limbs. Fewer than half responded to somatosensory stimulation. The neck region was the body area most effective in influencing unitary activity. The response pattern most often noted was an increase and decrease in discharge frequency when the body was moved towards and away from the recording electrode respectively. Change in discharge rate was observed to be primarily dependant upon neck velocity and not upon absolute neck position. Half of the somato-sensory units received input from either the forelimbs or the hindlimbs, while the remaining half responded to both.     

Non-cerebellar visual afferents to the vestibular nuclei involving the prepositus hypoglossal complex: An autoradiographic study in the rat

Summary Radioactive amino-acids were injected into the nucleus reticularis tegmenti pontis (NRTP) and the pretectum (PT) in the rat. Beside the labeling of the several nuclei which are known to receive afferents of either the NRTP and/or the PT, monosynaptic projections from these two structures to the prepositus hypoglossal complex (PHN) were demonstrated. Pretectal visual inputs to the vestibular nuclei (VN) may thus be conveyed not only by the classical PT-inferior olive-cerebellar route, but also by two other non-cerebellar ones involving the strong efferent projections of the PHN onto the VN. These last two pathways are strong candidates to account for the residual visual sensitivity of VN neurons after cerebellectomy or inferior olive lesions.Supported by CNRS (A.T.P. 8115)     

Influence of combined visual and vestibular cues on human perception and control of horizontal rotation

Summary Measurements are made of manual control performance in the closed-loop task of nulling perceived self-rotation velocity about an earth-vertical axis. Self-velocity estimation is modeled as a function of the simultaneous presentation of vestibular and peripheral visual field motion cues. Based on measured low-frequency operator behavior in three visual field environments, a parallel channel linear model is proposed which has separate visual and vestibular pathways summing in a complementary manner. A dual-input describing function analysis supports the complementary model; vestibular cues dominate sensation at higher frequencies. The describing function model is extended by the proposal of a non-linear cue conflict model, in which cue weighting depends on the level of agreement between visual and vestibular cues.Research supported in part by NASA Grants NSG 2032 and 2230. GLZ supported by an NIH National Research Service Award. GLZ currently at Bolt Beranek and Newman, Inc., Cambridge, MA, USA     

Senescence of human visual-vestibular interactions: smooth pursuit,optokinetic, and vestibular control of eye movements with aging

Natural aging entails progressive deterioration in a variety of biological systems. This study focuses on visual and vestibular influences on human eye movements as a function of aging. Eye movements were recorded (search-coil technique) during visual, vestibular, and combined stimuli in subjects across a broad range of ages (18–89 years). Two types of visual following were assessed: smooth pursuit (SP) of a small discrete target, and optokinetic (OKR) following of a large-field striped image. The vestibulo-ocular reflex (VOR) was studied during head rotation in darkness. Visualvestibular interactions were recorded during rotation in two ways: when the optokinetic scene was earth-fixed, resulting in visual enhancement of the VOR (VVOR), and when the visual image was head-fixed, allowing visual suppression of the VOR (VSVOR). Stimuli consisted of horizontal sinusoidal oscillations over the frequency range 0.025–4 Hz. Trials were analyzed to yield response gain (peak horizontal eye/stimulus velocities) and phase (asynchrony, in degrees, between eye and stimulus velocity signals). VOR gain in young subjects was greatest (near 0.9) at 2.5–4 Hz but declined steadily with decreasing frequency, while phase hovered near zero until 0.1 Hz and then developed a progressively increasing lead. Effects of advancing age were small, given the modest head velocities presented, and were most noticeable as an increase in phase lead and decline in gain at the lowest frequencies (0.1 Hz). The two forms of visual following and all conditions of visual-vestibular interactions displayed more prominent age-dependent changes. OKR and SP response characteristics (0.25–4 Hz) closely resembled each other. Gain was greatest at 0.25 Hz, while phase was near 0°. As frequency increased, gain declined while phase lag rose. However, both gain and phase lag tended to be slightly greater for OKR than for SP responses. Both SP and OKR response properties deteriorated progressively with increasing age, as witnessed by a progressive decline in gain and increase in phase lag, even at modest frequencies (e.g., 0.25–1.0 Hz). VVOR responses were generally closer to the ideal of 1.0 in gain and 0° in phase than either the VOR or visual following alone. A subtle but significant age-dependent decline in VVOR performance occurred at the lowest frequencies. VSVOR response characteristics were close to those of the VOR and VVOR at 4 Hz, where visual influences on eye movements are generally inconsequential. As frequency declined, visual suppression became more robust and gain dropped. The SP stimulus seemed surprisingly more effective than the OK scene in suppressing the VOR, but this effect is predicted by a linear model of visual-vestibular interactions. As age increased, visual influences on the VOR became progressively weaker, in concert with deterioration of visual following. The subjective sensation of circular vection (CV), a psychophysical measure of VVI, was assessed during optokinetic stimulation at 0.025 Hz. Interestingly, the likelihood and intensity of CV increased with aging, suggesting that visual inputs to the perception of self-motion are enhanced in the elderly. This may represent a form of visual compensation for age-dependent loss of vestibular self-rotation cues. In brief, the VOR, visual following, and their interactions display specific changes in response properties as a function of natural aging. The modifications may be interpreted as age-dependent deteriorations in the performance of systems underlying the control of human eye movements.     

Firing characteristics of vestibular nuclei neurons in the alert monkey after bilateral vestibular neurectomy

Summary After destruction of the peripheral vestibular system which is not activated by moving large-field visual stimulation, not only labyrinthine-ocular reflexes but also optokinetic-ocular responses related to the velocity storage mechanism are abolished. In the normal monkey optokinetic-ocular responses are reflected in sustained activity changes of central vestibular neurons within the vestibular nuclei. To account for the loss of optokinetic responses after labyrinthectomy, inactivation of central vestibular neurons consequent on the loss of primary vestibular activity is assumed to be of major importance. To test this hypothesis we recorded the neural activity within the vestibular nuclear complex in two chronically prepared Rhesus monkeys during a period from one up to 9 and 12 months after both vestibular nerves had been cut. The discharge characteristics of 829 cells were studied in relation to eye fixation, and to a moving small and large (optokinetic) visual stimulus producing smooth pursuit (SP) eye movements and optokinetic nystagmus (OKN). Units were grouped into different subclasses.After chronic bilateral vestibular neurectomy (BVN) we have found: (1) a rich variety of spontaneously active cells within the vestibular nuclear complex, which — as far as comparison before and after BVN is possible — belong to all subclasses of neurons functionally defined in normal monkey; and (2) no sustained activity changes which are related to the activation of the velocity storage mechanism; this is especially true for pure-vestibular, vestibular-pause and tonic-vestibular-pause cells in normal monkey which show a pure, pause and tonic-pause firing pattern after BVN. Neurons which are modulated by eye position are, however, modulated with the velocity of slow eye movements with comparable sensitivity during SP and OKN. Retinal slip is extremely rarely encoded. The results of the present study do not directly answer the question why the velocity storage mechanism is abolished after BVN but they suggest that only a small number of central vestibular cells may be inactivated by neurectomy.Supported by SNF grant no. 3.510-0.86     

Commissural and intrinsic connections of the vestibular nuclei in the rabbit: a retrograde labeling study

Summary The intrinsic and commissural projection of the vestibular nuclei were investigated by means of retrograde transport of normal (HRP) and wheatgerm-agglutinated horseradish peroxidase (WGA-HRP). It was found that within each vestibular complex, the superior (SV), medial (MV) and descending (DV) vestibular nuclei are reciprocally connected. A rostrocaudally oriented column of medium-sized and large neurons, comprising the central SV and the magnocellular MV (MVmc) receives input from the surrounding neurons and does not reciprocate this projection. Efferents from group y terminate in the SV, MV and DV. The infracerebellar nucleus (INF) as well as the interstitial nucleus of the VIII the nerve (IN) supply fibers to the MV and DV. The neurons that participate in the commissural projection are distributed throughout the vestibular complex with the exception of the lateral vestibular nucleus (LV) and group x. The largest number of cells was found in the MV. The HRP labeled cells show a tendency to cluster into rostrocaudally oriented groups. Each nucleus projects to more than one contralateral nucleus. Group y shows a more extensive contralateral projection than the bordering INF. It was concluded that quantitative differences in connectivity were present between a core region in the vestibular complex and peripheral parts. This core region comprises the central SV, the LV, the MVmc and extends into the rostral DV. It receives predominantly intrinsic input from the surrounding vestibular neurons and is in contrast to these latter neurons only minimally involved in the commissural projection.Abbreviations AChE acetylcholinesterase - bc brachium conjunctivum - bp brachium pontis - CE nucleus cuneatus externus - CO nuclei cochlearis - cr corpus restiforme - DV nucleus vestibularis descendens - DX nucleus dorsalis vagi - F nucleus fastigii - flm fasciculus longitudinalis medialis - gVII genu of the nervus facialis - group x, y, f groups x, y and f of Brodal - HRP horseradish peroxidase - IA nucleus interpositus anterior - IN nucleus interstitialis of nVIII - INF nucleus infracerebellaris - L nucleus lateralis - LV nucleus vestibularis lateralis - flm fasciculus longitudinalis medialis - MV nucleus vestibularis medialis - MVc caudal MV - MVmc magnocellular MV - MVpc parvocellular MV - nV nervus trigeminus - nVI nervus abducens - nVII nervus facialis - NV par nucleus vestibularis parabrachialis - PH nucleus prepositus hypoglossi - rV ramus descendens of nV - S nucleus and tractus solitarius - sad stria acustica dorsalis - SV nucleus vestibularis superior - tu tractus uncinatus - VI nucleus abducens - VM nucleus masticatorius - VOR vestibulo-ocular reflex - VP nucleus princeps trigemini - WGA-HRP wheatgerm-agglutinated HRP - XII nucleus hypoglossus     

Purkinje cell activity in the flocculus of vestibular neurectomized and normal monkeys during optokinetic nystagmus (OKN) and smooth pursuit eye movements

Summary 1. Single unit activity was recorded in the primate flocculus after the vestibular nerves were cut (bilateral vestibular neurectomy) during optokinetic nystagmus (OKN), smooth pursuit eye movements (SP) and whole field visual stimulation with gaze fixed on a stationary target light (OKN-suppression). Following vestibular neurectomy monkeys had no vestibular responses and no optokinetic after-nystagmus (OKAN) in the horizontal plane. However, OKN slow phases still reached steady state velocities of up to 100 deg/s. 2. After neurectomy, simple spike (SS) activity of Purkinje cells (P-cells) was modulated in relation to eye velocity, regardless of whether eye velocity was induced by a small target light moving in darkness (SP) or by a moving visual surround (OKN). In over 90% of the P-cells firing rate increased with eye velocity to the ipsilateral side and decreased with velocities to the contralateral side. Modulation in firing rate increased monotonically with increasing eye velocity. The strength of modulation was similar during SP and OKN for the same eye velocity. 3. The change in firing rate of P-cells in response to a sudden change in optokinetic stimulus velocity contained a component related to eye velocity and a component related to eye acceleration. Only a few P-cells were also modulated with image slip velocity during OKN-suppression. 4. The modulation of P-cells during SP and OKN was compared in normal and vestibular neurectomized monkeys. The sensitivity of floccular P-cells to eye velocity during SP was 1.14 imp·s^–1/deg·s^–1 in normal monkey and 1.28 imp·s^–1/deg·s^–1 after neurectomy. The similarity of eye velocity sensitivities demonstrates that neurectomy does not change the characteristics of floccular P-cell modulation during SP. In contrast, during OKN modulation of P-cells is quite different in normal and neurectomized monkey. In normal monkey, P-cells are modulated during steady state OKN for eye velocities above 40–60 deg/s only. This threshold velocity corresponds approximately to the maximal initial OKAN velocity (i.e. OKAN saturation velocity). After neurectomy, the threshold velocity is 0 deg/s and P-cells are modulated during steady state OKN also over ranges of eye velocities that do not cause a response in normal monkey. Sensitivities of P-cells to eye velocity during OKN for eye velocities above the threshold velocity are 1.0 imp·s^–1/deg·s^–1 in neurectomized monkey and 1.43 imp·s^–1/deg·s^–1 in normal monkey. 5. The hypothesis has been put forward that OKN slow phase velocity in normal monkey has two dynamically different components, a fast and a slow component. The results strongly suggest that the two components depend on different neuronal populations. Firing rate of floccular P-cells is modulated in relation to the fast component only. The results furthermore support the idea that it is the smooth pursuit system which may generate the fast component in the OKN slow phase velocity response.Supported by Swiss National Foundation for Scientific Research (Nr. 3.718-0.80 and 3.593-0.84)