Fragmental regulation of vestibuloocular responses

Intact pigeons were exposes to whole body centric and eccentric horizontal rotations in darkness during angular velocity trapezoids. The overall nystagmus alteration patterns were analysed. In 10 pigeons, all nystagmus alterations may be explained on the basis of the dynamics of peripheral otolith activity and central effects that are the same for all combinations of interacting inputs (type 1 patterns), whereas in other pigeons part of the nystagmus alterations were connected with some central effects that were individually specific and touch upon the responses to separate combinations of interacting input (type 2 patterns). It was observed the transformation of type 2 to type 1-patterns during the reiterated rotational trails and light nembutal anesthesia. A fragmentary control of the vestibulo-ocular responses seems to exist. This implies that the CNS is able to discern numerous kinds of bilaterally organized interacting inputs arising during different otolith membrane shifts.     

Habituation of horizontal eye nystagmus in pigeon during alternation of central and eccentric rotations

Intact pigeons were rotated in the horizontal plane in the dark in different positions relatively to the rotation axis. At central rotations, the pigeon's head was in the rotation axis whereas at eccentric rotations it was displaced from the axis. Series of central and eccentric rotations were alternated. Each series consisted of 2-5 rotations using angular velocity trapezoids. All stimuli producing habituation were used at most 14 times each. Eccentric rotations did not prevent a gradual decrease of peak velocities of the slow component of primary nystagmus on transition from one series of central rotations to another in 17 pigeons (group 1). The increase of peak velocities was observed in 2 pigeons (group 2). In group 1, a direct dependence among alterations of this parameter of primary nystagmus, modifications of its duration, and variations of peak velocities of secondary nystagmus, were observed. If two identical stimuli did not follow in sequence directly, the effect of the second one produced same nystagmus changes as were observed in present pigeon by comparison of the first and last responses in the series of the central rotations. If they follow one by one, in many cases the second stimulus in the pair produced an increase of peak velocity of primary and secondary nystagmus and rise of delay of the point of primary nustagmus peak velocity. These variations were not random (probability, > 95%).     

The interstitial nucleus of Cajal of the cat. I. Neurons activity related to vertical eye movements

Interstitial nucleus of Cajal (INC) neurons activity was studied during vertical optokinetic nystagmus (OKN) and after-nystagmus (OKAN) in awake cats lying on their right side. The activity of one hundred neurons was recorded in the left INC and analysed in relation with the vertical component of OKN and OKAN. The activity of 27 neurons was correlated either to eye position or to both eye velocity and eye position; 18 of these neurons were recorded in their on-direction and their off-direction. The analysis of the 18 neurons showed that the activity of 8 of them was correlated to eye position in the on-direction and in the off-direction and the correlation to eye position was higher than to eye velocity; these neurons are considered as position neurons. Seven other neurons had a higher correlation to eye position that to eye velocity in the on-direction and this relation reversed in the off-direction, these neurons are considered as position-velocity neurons. Thirty two burst-neurons were activated only during quick phases of OKN and OKAN and they were silent during slow phases and periods of fixation. Nine burst neurons had an upward on-direction and 23 neurons a downward on-direction. The eye velocity-average burst frequency (ABF) and quick phase duration-burst duration relationships had low correlations and suggested that INC burst neurons were excitatory premotor neurons. Statistical analysis showed that downward on-direction burst neurons had a higher ABF that upward on-direction burst neurons. Moreover, during OKN and OKAN, the velocity sensitivity of INC burst neurons was the same. The activity of the remaining neurons (41 neurons) was not quantitatively correlated to vertical and horizontal eye movements; they were classified as irregular tonic neurons. This study shows that INC neurons carry an eye position signal which was never reported before. This supports the results of INC lesion studies which showed that INC is involved in the vertical velocity to position integration. Moreover, there is an up versus down asymmetry in the frequency of INC burst neurons.     

Experimental research on maculo-ampullar interactions in normal subjects

The AA. investigated 20 normal subjects to evaluated the influence of the otolith organ on the nystagmus, induced by angular accelerations in centrifugal and eccentric rotations. The nystagmus has been recorded by electronystagmostical analysis. The results show that centrifugal and eccentric tests reduce the nystagmus intensity and particularly the amplitude, duration and slow-phase velocity during the per-rotatory accelerations. These findings confirm that the macular activity may inhibit the nystagmus reactions induced by angular acceleration.     

A model of the nystagmus induced by off vertical axis rotation

A three-dimensional model is proposed that accounts for a number of phenomena attributed to the otoliths. It is constructed by extending and modifying a model of vestibular velocity storage. It is proposed that the otolith information about the orientation of the head to gravity changes the time constant of vestibular responses by modulating the gain of the velocity storage feedback loop. It is further proposed that the otolith signals, such as those that generate L-nystagmus (linear acceleration induced nystagmus), are partially coupled to the vestibular system via the velocity storage integrator. The combination of these two hypotheses suggests that a vestibular neural mechanism exists that performs correlation in the mathematical sense which is multiplication followed by integration. The multiplication is performed by the otolith modulation of the velocity storage feedback loop gain and the integration is performed by the velocity storage mechanism itself. Correlation allows calculation of the degree to which two signals are related and in this context provides a simple method of determining head angular velocity from the components of linear acceleration induced by off-vertical axis rotation. Correlation accounts for the otolith supplementation of the VOR and the sustained nystagmus generated by off-vertical axis rotation. The model also predicts the cross-coupling of horizontal and vertical optokinetic afternystagmus that occurs in head-lateral positions and the reported effects of tilt on vestibular responses.     

Three dimensional eye movement analysis during off vertical axis rotation in human subjects

Three dimensional analysis of eye movements during OVAR was performed in 37 healthy human subjects using the computerized image recognition technique developed by us. The modulation component of eye movement was observed in all three components (horizontal, vertical and torsional), whereas the bias component was only clearly seen in the horizontal eye movement. The phase lag of the torsional component was quite consistent with a small variation between each subject with respect to the head position. The phase of vertical eye movement was, however, less consistent compared to that of the horizontal and torsional eye movements. From these results, in human subjects, there should be some differences in the dynamic function of the otolith system compared to that observed in monkeys.     

Automatic detection and removal of fast phases from nystagmographic recordings by optimal thresholding

Recording of ocular nystagmus during vestibular tests does not measure the true response of the vestibulo-ocular reflex (VOR), because the VOR response (so-called slow phase of nystagmus) is interrupted by resetting saccades (so-called fast phase of nystagmus). In order to extract the real VOR contribution, saccades must be removed. In most of the nystagmus processing algorithms, saccade removal requires a human operator to choose a suitable eye velocity threshold able to separate fast from slow nystagmus phases. In the present report a fully automatic removal system is presented which selects an optimal velocity threshold by computing the VOR frequency response and maximizing its coherence function.     

Oculomotor areas in the rabbit's midbrain and pretectum

The role of some meso- and diencephalic structures in eye movements was investigated by ablation and stimulation experiments. Optokinetic nystagmus was abolished by small lesions in the lateral pretectum, but not by complete removal of the superior colliculi. Stimulation of the superior colliculus and other visual centers was effective in eliciting nystagmus (slow phase ipsilateral), but the most efficient trigger zones are found in the lateral pretectum and the midbrain tegmentum. Only from these areas could nystagmus still be elicited after degeneration of the primary optic fibers. The lateral pretectal trigger zone is probably identical with the nucleus of the optic tract. It is postulated that this nucleus is an essential station for horizontal optokinetic reactions. Saccades were obtained by stimulation of the mesencephalic central grey, but not for any visual centers such as the superior colliculus.     

Oculomotor areas in the rabbits midbrain and pretectum.

The role of some meso- and diencephalic structures in eye movements was investigated by ablation and stimulation experiments. Optokinetic nystagmus was abolished by small lesions in the lateral pretectum, but not by complete removal of the superior colliculi. Stimulation of the superior colliculus and other visual centers was effective in eliciting nystagmus (slow phase ipsilateral), but the most efficient trigger zones are found in the lateral pretectum and the midbrain tegmentum. Only from these areas could nystagmus still be elicited after degeneration of the primary optic fibers. The lateral pretectal trigger zone is probably identical with the nucleus of the optic tract. It is postulated that this nucleus is an essential station for horizontal optokinetic reactions. Saccades were obtained by stimulation of the mesencephalic central grey, but not for any visual centers such as the superior colliculus.     

Alterations in rat horizontal vestibulo-ocular reflex phase as a function of orientation in gravity

The effect of changes in static and dynamic gravity signals on the phase accuracy of the horizontal vestibulo-ocular reflex (HVOR) was studied in rats using chronically implanted scleral search coils to monitor eye movements. Rats were sinusoidally rotated using a range of different frequencies (0.035-2 Hz) in a plane which always activated the horizontal semicircular canals but in one of three different orientations with regard to gravity which differentially activated the otolith organs: 1) upright-normal static gravity signal, no dynamic otolith activation; 2) inverted-inverted static gravity signal, no dynamic otolith activation; 3) on-side-dynamic activation of the otolith organs. In the upright orientation, the HVOR shows a phase advance at 0.2 Hz and below but not at 0.5 Hz and above. Phase accuracy of the HVOR was further degraded in the inverted orientation with rats showing large phase leads at 0.2 Hz and below. In contrast, accuracy of the HVOR was significantly improved at 0.2 Hz and below in the on-side orientation with phase accurate eye movements down to the lowest frequency tested. The results further support the idea that otolith organs play an important role in VOR generation by supplementing the semicircular canals' response to angular head movements.     

Effect of active head movements about the pitch, roll, and yaw axes on human optokinetic afternystagmus

Effects of active head movements about the pitch, roll, or yaw axes on horizontal optokinetic afternystagmas (OKAN) were examined in 16 subjects to test the hypothesis that otolith organ mediated activity induced by a change in head position can couple to the horizontal velocity storage in humans. Active head movements about the pitch axis, forwards or backwards, produced significant OKAN suppression. Pitch forward head movements exerted the strongest effect. Active head movements about the roll axis towards the right also produced OKAN suppression but only if the tilted position was sustained. No suppression was observed following sustained yaw. However, an unsustained yaw left movement after rightward drum rotation significantly enhanced OKAN. Sustained head movement trials did not significantly alter subsequent control trials. In contrast, unsustained movements about the pitch axis, which involve more complex interactions, exerted long-term effects on subsequent control trials. We conclude that otolith organ mediated activity arising from pitch or roll head movements couples to the horizontal velocity storage in humans, thereby suppressing ongoing OKAN. Activity arising from the horizontal canals during an unsustained yaw movement (observed mainly with yaw left), following drum rotation in a direction contralateral to the movement, may also couple to the velocity storage, resulting in increased activity instead of suppression.     

Static cervical-ocular reflex and optokinetic reflex interaction in rabbits

Summary The effect that tonic eye deviations, induced by angular deviation of the torso, have on the characteristics of optokinetic (OK) nystagmus was studied in rabbits. When the slow component of the OK nystagmus moved in the direction of the tonic eye deviation, the amplitude of the slow and fast components of the nystagmus was decreased and their frequency was increased, whereas when the slow component moved in the opposite direction, the amplitude and the frequency of the nystagmus were not different from those when the head and torso were aligned.Under the influence of neck reflexes, the total range of eye movements was double that when the torso was aligned with the head. The place in the orbit where the fast-component is initiated — the so-called fast-component threshold — was deviated in the direction of the neck-reflex-induced tonic eye deviation. The characteristics of the fast component, however, except for its amplitude, were not affected by the change of location of the fast-component threshold.These data indicate that the OK reflex function, as judged by measurement of the slow component velocity, is not affected by neck-vestibular reflexes. They also show that the fast-component threshold is dependent on parameters other than the actual orbital position and that there must be an internal representation of the range of possible eye movements within the brain to regulate the production of fast components.Abbreviations OK optokinetic - CW clockwise - CCW counterclockwise - CNS central nervous system This work was supported by grants NS07059, NS09823, and NS08335 from the National Institutes of Health     

The role of background movement in goldfish vision

In experiments described in the literature objects presented to restrained goldfish failed to induce eye movements like fixation and/or tracking. We show here that eye movements can be induced only if the background (visual surround) is not stationary relative to the fish but moving. We investigated the influence of background motion on eye movements in the range of angular velocities of 5–20° s⁻¹. The response to presentation of an object is a transient shift in mean horizontal eye position which lasts for some 10 s. If an object is presented in front of the fish the eyes move in a direction such that it is seen more or less symmetrically by both eyes. If it is presented at ±70° from the fish's long axis the eye on the side of the object moves in the direction that the object falls more centrally on its retina. During these object induced eye responses the typical optokinetic nystagmus of amplitude of some 5° with alternating fast and slow phases is maintained, and the eye velocity during the slow phase is not modified by presentation of the object. Presenting an object in front of stationary or moving backgrounds leads to transient suppression of respiration which shows habituation to repeated object presentations. Accepted: 14 April 2000     

运动图形刺激时家兔的视动震颤反应

旨在用实验方法研究家兔的视动震颤（ＯＫＮ）眼动特点以及单侧前庭迷路损伤对ＯＫＮ的影响,结果表明：单眼刺激时,家兔的ＯＫＮ反应存在着从颞侧到鼻侧方向的方向优势;恒定速度刺激时,刺激开始后,家兔的ＯＫＮ眼动跟踪速度具有从小到大最后趋于稳态的建立过程,刺激消失后,存在眼动速度由大到小直到消失的视动后震颤（ＯＫＡＮ）反应,这两个过程反应了ＯＫＮ系统中可能存在速度存储机制及其对ＯＫＮ眼动的控制作用;单侧前庭     

A linear canal-otolith interaction model to describe the human vestibulo-ocular reflex

A control systems model of the vestibulo-ocular reflex (VOR) originally derived for yaw rotation about an eccentric axis (Crane et al. 1997) was applied to data collected during ambulation and dynamic posturography. The model incorporates a linear summation of an otolith response due to head translation scaled by target distance, adding to a semi-circular canal response that depends only on angular head rotation. The results of the model were compared with human experimental data by supplying head angular velocity as determined by magnetic search coil recording as the input for the canal branch of the model and supplying linear acceleration as determined by flux gate magnetometer measurements of otolith position. The model was fit to data by determining otolith weighting that enabled the model to best fit the data. We fit to the model experimental data from normal subjects who were: standing quietly, walking, running, or making active sinusoidal head movements. We also fit data obtained during dynamic posturography tasks of: standing on a platform sliding in a horizontal plane at 0.2 Hz, standing directly on a platform tilting at 0.1 Hz, and standing on the tilting platform buffered by a 5-cm thick foam rubber cushion. Each task was done with the subject attending a target approximately 500, 100, or 50 cm distant, both in light and darkness. The model accurately predicted the observed VOR response during each test. Greater otolith weighting was required for near targets for nearly all activities, consistent with weights for the otolith component found in previous studies employing imposed rotations. The only exceptions were for vertical axis motion during standing, sliding, and tilting when the platform was buffered with foam rubber. In the horizontal axis, the model always fit near target data better with a higher otolith component. Otolith weights were similar with the target visible and in darkness. The model predicts eye movement during both passive whole-body rotation and free head movement in space implying that the VOR is controlled by a similar mechanism during both situations. Factors such as vision, proprioception, and efference copy that are available during head free motion but not during whole-body rotation are probably not important to gaze stabilization during ambulation and postural stabilizing movement. The linearity of the canal-otolith interaction was tested by re-analysis of the whole body rotation data on which the model is based (Crane et al. 1997). Normalized otolith-mediated gain enhancement was determined for each axis of rotation. This analysis uncovered minor non-linearities in the canal-otolith interaction at frequencies above 1.6 Hz and when the axis of rotation was posterior to the head. Received: 11 March 1998 / Received in revised form: 1 March 1999     

Development of Eye Position Dependency of Slow Phase Velocity during Caloric Stimulation

The nystagmus in patients with vestibular disorders often has an eye position dependency, called Alexander’s law, where the slow phase velocity is higher with gaze in the fast phase direction compared with gaze in the slow phase direction. Alexander’s law has been hypothesized to arise either due to adaptive changes in the velocity-to-position neural integrator, or as a consequence of processing of the vestibular-ocular reflex. We tested whether Alexander’s law arises only as a consequence of non-physiologic vestibular stimulation. We measured the time course of the development of Alexander’s law in healthy humans with nystagmus caused by three types of caloric vestibular stimulation: cold (unilateral inhibition), warm (unilateral excitation), and simultaneous bilateral bithermal (one side cold, the other warm) stimulation, mimicking the normal push-pull pattern of vestibular stimulation. Alexander’s law, measured as a negative slope of the velocity versus position curve, was observed in all conditions. A reversed pattern of eye position dependency (positive slope) was found <10% of the time. The slope often changed with nystagmus velocity (cross-correlation of nystagmus speed and slope was significant in 50% of cases), and the average lag of the slope with the speed was not significantly different from zero. Our results do not support the hypothesis that Alexander’s law can only be observed with non-physiologic vestibular stimulation. Further, the rapid development of Alexander’s law, while possible for an adaptive mechanism, is nonetheless quite fast compared to most other ocular motor adaptations. These results suggest that Alexander’s law may not be a consequence of a true adaptive mechanism.     

Three-dimensional head angular velocity detection from otolith afferent signals

Afferent signals from the otolith organs can produce compensatory eye position and velocity signals which has been described as linear vestibulo-ocular reflex (LVOR). The afferent otolith signals carry information about head orientation and changes of head orientation relative to gravity. A head orientation (tilt) related position signal can be obtained from population vector coding of tonic otolith afferent signals during static or dynamic head tilts, which in turn could produce compensatory eye position signals in the LVOR. On the other hand, eye angular velocity signals may be extracted, as proposed in this study, from the population response of tilt-velocity sensitive otolith afferents. Such afferents are shown to encode instantaneous head orientation relative to gravity at onset of a head movement and, as the movement continues, the projection of head angular velocity onto the earth-horizontal plane, indicating the instantaneous direction of movement relative to gravity. Angular velocity components along the earth-vertical direction which are not directly encoded by otolith afferents can be detected by central signal processing. Central reconstruction of 3D head angular velocity allows to obtain information about absolute head orientation in space even in the absence of semicircular canal related information. Such information is important for generating compensatory eye movements as well as for dynamic control of posture.     

A random walk model of fast-phase timing during optokinetic nystagmus

 Most vertebrate animals produce optokinetic nystagmus in response to rotation of their visual surround. Nystagmus consists of an alternation of slow-phase eye rotations, which follow the surround, and fast-phase eye rotations, which quickly reset eye position. The time intervals between fast phases vary stochastically, even during optokinetic nystagmus produced by constant velocity rotation of a uniform surround. The inter-fast-phase interval distribution has a long tail, and intervals that are long relative to the mode become even more likely as constant surround velocity is decreased. This paper provides insight into fast-phase timing by showing that the process of fast-phase generation during constant velocity optokinetic nystagmus is analogous to a random walk with drift toward a threshold. Neurophysiologically, the output of vestibular nucleus neurons, which drive the slow phase, would approximate a random walk with drift because they integrate the noisy, constant surround velocity signal they receive from the visual system. Burst neurons, which fire a burst to drive the fast phase and reset the slow phase, are brought to threshold by the vestibular nucleus neurons. Such a nystagmic process produces stochastically varying inter-fast-phase intervals, and long intervals emerge naturally because, as drift rate (related to surround velocity) decreases, it becomes more likely that any random walk can meander for a long time before it crosses the threshold. The theoretical probability density function of the first threshold crossing times of random walks with drift is known to be that of an inverse Gaussian distribution. This probability density function describes well the distributions of the intervals between fast phases that were either determined experimentally, or simulated using a neurophysiologically plausible neural network model of fast-phase generation, during constant velocity optokinetic nystagmus. Received: 1 June 1995/Accepted in revised form: 15 February 1996     

Convergence rotatory nystagmus under unusual conditions of semicircular canal and otolith stimulation

The characteristics of interaction between two vestibular subsystems (otiliths and semicircular canals) were studied by means of binocular (bilateral) videooculographic recording of eye movements in 43 men aged from 19 to 41 years that had been found healthy upon aviation physical examination. The time course of horizontal vestibular nystagmus was analyzed separately for each eye in subjects who bent forward and straightened up in the sagittal plane while being rotated about the vertical body axis in an electrically driven rotating chair. This combined rotation caused interocular asymmetric nystagmus in 91% of the subjects and convergence rotatory nystagmus in 42% of the subjects. A hypothesis on the mechanism of interocular asymmetric nystagmus caused by the combined rotation and convergence rotatory nystagmus as its special case has been advanced. The hypothesis allows for independent nystagmic mechanisms (subsystems) for the right and left eyes.