Johannes van der Steen

Abstract: Human orienting behaviour requires a complex interaction between the visual and the oculomotor system. We present orienting gaze data measured in children using a remote eye tracking system. The aim of the study was to validate a data analysis method which did not require off-line correction of data gaps due to eye blinking or inadequate gaze tracking. In two sessions, blocks of short movies, amongst others cartoons, were shown to 35 children (2-9 years) for a test-retest analysis. The cartoons were subsequently shown in one of the monitor corners. Orienting eye movements were analysed on the basis of saccadic reaction time (SRT), reaction time to fixation (RTF) of cartoon and gaze fixation area (GFA) Differences were tested for significance using the Wilcoxon-signed ranks test and reliability was assessed using the intraclass correlation coefficient (ICC). SRT values could be calculated in approximately 50% of gaze data and ranged between 150(30)ms (mean(SD)) and 390(190)ms (average SRT(min) and SRT(max) values of all subjects). RTF values could be derived in approximately 90% of gaze data with an average RTF(min) of 210(30)ms and RTF(max) of 570(160)ms. Test-retest analysis showed a significant increase of GFA during the second session with approximately 5% (P<0.05). The reliability of RTF(min) and GFA was best with an ICC of 0.84 and 0.80, respectively (P<0.0001). We conclude that remote eye tracking is well suited for quantification of timing and executing oculomotor fixations during orienting behaviour tasks. The presented method may be applied in young children with developmental disorders or brain damage.

Abstract: The three-dimensional vestibulo-ocular reflex (3D VOR) ideally generates compensatory ocular rotations not only with a magnitude equal and opposite to the head rotation but also about an axis that is collinear with the head rotation axis. Vestibulo-ocular responses only partially fulfill this ideal behavior. Because animal studies have shown that vestibular stimulation about particular axes may lead to suboptimal compensatory responses, we investigated in healthy subjects the peaks and troughs in 3D VOR stabilization in terms of gain and alignment of the 3D vestibulo-ocular response. Six healthy upright sitting subjects underwent whole body small amplitude sinusoidal and constant acceleration transients delivered by a six-degree-of-freedom motion platform. Subjects were oscillated about the vertical axis and about axes in the horizontal plane varying between roll and pitch at increments of 22.5 degrees in azimuth. Transients were delivered in yaw, roll, and pitch and in the vertical canal planes. Eye movements were recorded in with 3D search coils. Eye coil signals were converted to rotation vectors, from which we calculated gain and misalignment. During horizontal axis stimulation, systematic deviations were found. In the light, misalignment of the 3D VOR had a maximum misalignment at about 45 degrees . These deviations in misalignment can be explained by vector summation of the eye rotation components with a low gain for torsion and high gain for vertical. In the dark and in response to transients, gain of all components had lower values. Misalignment in darkness and for transients had different peaks and troughs than in the light: its minimum was during pitch axis stimulation and its maximum during roll axis stimulation. We show that the relatively large misalignment for roll in darkness is due to a horizontal eye movement component that is only present in darkness. In combination with the relatively low torsion gain, this horizontal component has a relative large effect on the alignment of the eye rotation axis with respect to the head rotation axis.

Abstract: Physical whole-body vibration (WBV) exercises become available at various levels of intensity. In a first series of measurements, we investigated 3-dimensional platform accelerations of three different WBV devices without and with three volunteers of different weight (62, 81 and 100 kg) in squat position (150 degrees knee flexion). The devices tested were two professional devices, the PowerPlate and the Galileo-Fitness, and one home-use device, the PowerMaxx. In a second series of measurements, the transmission of vertical platform accelerations of each device to the lower limbs was tested in eight healthy volunteers in squat position (100 degrees knee flexion). The first series showed that the platforms of two professional devices vibrated in an almost perfect vertical sine wave at frequencies between 25-50 and 5-40 Hz, respectively. The platform accelerations were slightly influenced by body weight. The PowerMaxx platform mainly vibrated in the horizontal plane at frequencies between 22 and 32 Hz, with minimal accelerations in the vertical direction. The weight of the volunteers reduced the platform accelerations in the horizontal plane but amplified those in the vertical direction about eight times. The vertical accelerations were highest in the Galileo (approximately 15 units of g) and the PowerPlate (approximately 8 units of g) and lowest in the PowerMaxx (approximately 2 units of g). The second series showed that the transmission of vertical accelerations at a common preset vibration frequency of 25 Hz were largest in the ankle and that transmission of acceleration reduced approximately 10 times at the knee and hip. We conclude that large variation in 3-dimensional accelerations exist in commercially available devices. The results suggest that these differences in mechanical behaviour induce variations in transmissibility of vertical vibrations to the (lower) body.

Abstract: PURPOSE: This study compared the performance of a video-based infrared three-dimensional eye tracker device (Chronos) with the scleral search coil method. METHODS: Three-dimensional eye movements were measured simultaneously with both systems during fixation, saccades, optokinetic stimulation, and vestibular stimulation. RESULTS: Comparison of fixation positions between -15 degrees and +15 degrees showed that horizontal and vertical eye position signals of the two systems were highly correlated (R2 = 0.99). Torsion values measured by coils and the video system were significantly different (P < 0.001). Saccade main sequence parameters of coil and video signals were in good agreement. Gains of torsion in response to optokinetic stimulation (cycloversion and cyclovergence) were not significantly different (P > 0.05). Gain values of the vestibulo-ocular reflex as determined from coil and video signals showed good agreement for rotations. However, there was more variability in the video signals for translations, possibly due to relative motion between the head and cameras. CONCLUSIONS: Lower time resolution, possible instability of the head device of the video system, and inherent small instabilities of pupil tracking algorithms make the coil system the best choice when measuring eye movement responses with high precision or when involving high-frequency head motion. For less demanding and for static tests and measurements longer than a half an hour, the latest generation infrared video system is a good alternative to scleral search coils. However, the quality of torsion of the infrared video system is less compared with scleral search coils and needs further technological improvement.

Abstract: A new technique is introduced to measure linear and angular vestibulo-ocular responses in three dimensions. Using a three-dimensional motion platform, human subjects underwent whole-body rotations and translations. Eye movements were measured with an infrared video recording device and/or with scleral search coils. Subjects were tested with sinusoidal stimulation and impulses under light and dark conditions. The results show that for sinusoidal stimulation, torsion compensatory eye movements (roll stimulation) have a low gain compared to horizontal (yaw) and vertical (pitch) compensatory eye movements. With impulses, we reliably assessed gain and delay for rotations in yaw, pitch, and roll. Under this stimulus condition the gain for roll (torsion eye movements) was also low compared to yaw and pitch (horizontal and vertical eye movements). For translations, the gain of eye-movement responses varied between 0.7 and 1 in the light. In the dark, responses were lower and more variable.

Abstract: Superior-canal dehiscence syndrome (SCDS) was first described in 1998; it manifests as symptoms of dizziness and oscillopsia when a sound or pressure stimulus is given to the ear concerned. SCDS is caused by dehiscence ofpetrosal bone overlying the superior, vertical semicircular canal, creating a 'third mobile window' to the labyrinth, in addition to the oval and round windows. Characteristic for this syndrome are eye movements with a vertical-torsion direction, evoked by sound stimuli, or pressure changes via the external ear canal and the Valsalva manoeuvre. The anomaly of the bony structure is visible on a CT scan. With its specific symptoms and characteristics, SCDS is an abnormality of the vestibular system that is relatively easy to diagnose. The syndrome may be treated with conservative or surgical means.

Abstract: Cerebellar long-term depression (LTD) at parallel fibre-Purkinje cell (P-cell) synapses is thought to embody neuronal information storage for motor learning. Transgenic L7-protein kinase C inhibitor (PKCI) mice in which cerebellar LTD is selectively blocked do indeed exhibit impaired adaptation in the vestibulo-ocular reflex (VOR) while their default oculomotor performance is unaffected. Although supportive, these data do not definitively establish a causal link between memory storage required for motor learning and cerebellar LTD. As the L7-PKCI transgene is probably activated from the early stages of P-cell development, an alternative could be that P-cells develop abnormal signals in L7-PKCI mutants, disturbing mechanisms of motor learning that rely on proper P-cell outputs. To test this alternative hypothesis, we studied simple spike (SS) and complex spike (CS) activity of vertical axis P-cells in the flocculus of L7-PKCI mice and their wild-type littermates during sinusoidal optokinetic stimulation. Both SS and CS discharge dynamics appeared to be very similar in wild-type and transgenic P-cells at all stimulus frequencies (0.05-0.8 Hz). The CS activity of all vertical axis cells increased with contralateral stimulus rotation and lagged ipsiversive eye velocity by 165-180 degrees. The SS modulation was roughly reciprocal to the CS modulation and lagged ipsiversive eye velocity by approximately 15 degrees. The baseline SS and CS discharge characteristics were indistinguishable between the two genotypes. We conclude that the impaired VOR learning in L7-PKCI mutants does not reflect fundamental aberrations of the cerebellar circuitry. The data thus strengthen the evidence that cerebellar LTD is implicated in rapid VOR learning but not in the development of normal default response patterns.

Abstract: The flocculus of the rabbit is involved in the plasticity of compensatory eye movements. It is generally assumed that the climbing fiber input to floccular Purkinje cells encodes "retinal slip," which in turn would be a measure for the oculomotor performance error. To test this, we used transparent motion stimuli, creating a retinal slip signal that broke up this relation. We recorded the ensuing oculomotor behavior and complex spike activity of floccular Purkinje cells. Complex spike modulation in response to transparent stimulation was identical to that of a single optokinetic pattern, despite considerably different retinal slip. These results suggest that the climbing fiber code may be effectively related to the eye movement performance error, rather than to retinal slip.

Abstract: We studied the coordination of binocular eye movements in human subjects with alternating exotropia (divergent strabismus). Binocular saccades were recorded in six subjects during binocular and monocular viewing. Subjects were instructed to make saccades between two continuously lit targets (LED's) presented in an isovergence array (with the straight-ahead target 130 cm from the eyes) in a dimly lit room. For saccades up to 20 degrees amplitude, there were no large differences in the dynamics of the saccades between control and exotropic subjects. However, for larger amplitudes subjects frequently alternated the eye of fixation during saccades. That is, subjects fixated the left target with the left eye and the right target with the right eye. The alternation in eye fixation at the end of the saccade was taken into account in the programming of the saccades. The amplitudes of the alternating saccades were approximately equal to the target amplitude minus the strabismus angle. We conclude that for those saccades where alternation occurs, there is not only a change in the eye of fixation, but also a change in the target representation provided by either eye. Thus, in this group of strabismic patients, saccades may be programmed in a retina-centered coordinate system, if we assume that for making a saccade to a new target in the contralateral visual field its representation on the temporal retinal field of the currently fixating eye is suppressed and the retinotopic target information is derived from the non-fixating eye. In executing the saccade, the non-fixating eye automatically becomes the fixating eye.

Abstract: A longstanding but still controversial hypothesis is that long-term depression (LTD) of parallel fiber-Purkinje cell synapses in the cerebellum embodies part of the neuronal information storage required for associative motor learning. Transgenic mice in which LTD is blocked by Purkinje cell-specific inhibition of protein kinase C (PKC) (L7-PKCI mutants) do indeed show impaired adaptation of their vestibulo-ocular reflex, whereas the dynamics of their eye movement performance are unaffected. However, because L7-PKCI mutants have a persistent multiple climbing fiber innervation at least until 35 d of age and because the baseline discharge of the Purkinje cells in the L7-PKCI mutants is unknown, factors other than a blockage of LTD induction itself may underlie their impaired motor learning. We therefore investigated the spontaneous discharge of Purkinje cells in alert adult L7-PKCI mice as well as their multiple climbing fiber innervation beyond the age of 3 months. We found that the simple spike and complex spike-firing properties (such as mean firing rate, interspike interval, and spike count variability), oscillations, and climbing fiber pause in the L7-PKCI mutants were indistinguishable from those in their wild-type littermates. In addition, we found that multiple climbing fiber innervation does not occur in cerebellar slices obtained from 3- to 6-month-old mutants. These data indicate (1) that neither PKC inhibition nor the subsequent blockage of LTD induction disturbs the spontaneous discharge of Purkinje cells in alert mice, (2) that Purkinje cell-specific inhibition of PKC detains rather than prevents the developmental conversion from multiple to mono-innervation of Purkinje cells by climbing fibers, and (3) that as a consequence the impaired motor learning as observed in older adult L7-PKCI mutants cannot be attributable either to a disturbance in the baseline simple spike and complex spike activities of their Purkinje cells or to a persistent multiple climbing fiber innervation. We conclude that cerebellar LTD is probably one of the major mechanisms underlying motor learning, but that deficits in LTD induction and motor learning as observed in the L7-PKCI mutants may only be reflected in differences of the Purkinje cell signals during and/or directly after training.

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Abstract: The way a rabbit moves its eyes in response to a stimulus consisting of two moving random dot patterns largely depends on the relative luminances of the two patterns. Concurrent rotation of the animal enhances the response to the visual pattern that represents the same head movement as the vestibular stimulation. In this paper we investigate the role that the flocculus plays in this behaviour. We injected the non-selective acetylcholine agonist carbachol into the flocculus. These injections are known to increase the gain of the optokinetic reflex, but have a smaller effect on the vestibulo-ocular reflex. We investigated the effect on the oculomotor response to (vestibulo-) transparent stimuli, where one pattern oscillated sinusoidally and the other pattern was stable with respect to the head. We found that the injections caused a higher response gain at a lower luminance of the oscillating pattern. Furthermore the influence of concurrent vestibular stimulation decreased. These findings agree with a role of the flocculus that is downstream of the visual normalisation, but upstream of the visual-vestibular interaction.

Abstract: In the present study, eve movements during the copying of a pattern were analyzed to compare visual strategies of adults and children. Subjects had to build an accurate copy of spatial block patterns. Tested variables were the incidence and the duration of ocular dwelling in the pattern area (where the pattern to be copied was located), the work area (where the copy was made), and the source area (where the blocks were that could be used for creating the copy). Furthermore, to unravel employed strategies, sequences of dwelled areas were investigated. Previous studies reported that adults employ repetitive visual scanning strategies to accomplish the task instead of strategies depending upon an internal representation. The present results show that the II children, within the ages of 7 to 12 years, made more eye movements and fixations of longer duration during copy tasks than the 11 adults. The visual strategies of the children were highly comparable to those of adults. Memory was restricted to one block, while color and location seemed to be remembered together.

Abstract: Transparent motion is a visual stimulus condition that generates multiple motion vectors on the retina that can differ in speed, direction, and/or luminance. Transparent motion creates a conflict for retinal stabilization. In this study we investigated the effect of transparent visual motion on the oculomotor reflexes that provide retinal stabilization in the rabbit. In the first experimental condition, the animals were stationary. We presented one stationary and one oscillating visual pattern to the animals while varying the luminance of the patterns. We found that the optokinetic eye movement responses were fully determined by the luminance of the individual visual inputs, weighted for the total luminance. No effect of absolute stimulus intensity was found. In the second experimental condition we oscillated the animals, while using an identical visual stimulation paradigm. The contribution of the vestibulo-ocular reflex enhanced the response to the visual pattern, which was in agreement with the vestibular stimulus. This effect of vestibular stimulation was independent of the absolute intensity of the visual stimuli. From this result we conclude that the weighting process of the transparent visual patterns occurs upstream from the site of the visual-vestibular interaction. Both the visual weighting and the visual-vestibular interaction were dependent on stimulus frequency. In line with the properties of the visual and vestibular stabilization reflexes in isolation, the contribution of the vestibular system increased, whereas the influence of the optokinetic system decreased with increasing stimulus frequency.

Abstract: We studied gaze-shift dynamics during several gaze-shift tasks and during reading, in five subjects with convergence insufficiency (C.I., a diminished ability to converge), and in ten subjects without C.I. Furthermore, we studied the effect of vergence training in order to verify previous claims that orthoptic exercises can improve vergence performance. We recorded binocular eye movements with the scleral coil technique. Subjects switched fixation between nearby and distant light emitting diodes (LEDs) arranged in isovergence arrays (distances 35 and 130 cm) in a dimly lit room. In both the C.I. and non-C.I. group, two classes of subjects occurred: vergence responders and saccadic responders. During pure vergence tasks, saccadic responders made saccades with no or little vergence; vergence responders made vergence movements with no or small saccadic components. In saccadic responders, fixation of nearby targets was monocular. Subjects with a preferred eye, according to our determination, used the preferred eye. The five C.I. subjects showed idiosyncratic responses with insufficient vergence during most trials. They all had a tendency to alternate fixation between the left and right eye. Vergence-version tasks always elicited larger vergence components than pure vergence tasks. During a reading task, vergence angles were more accurate than during gaze-shifts between LEDs. After the pre-training sessions, nine subjects (one of which had C.I.) practised a pure vergence task three times a day for at least 2 weeks. Vergence amplitudes of four of these subjects were larger after training. We conclude that vergence training can change oculomotor performance. Although C.I. is often associated with abnormal vergence dynamics, there are no typical C.I. vergence dynamics. Unstable monocular preferences may play a role in the aetiology of C.I.

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Abstract: We attempted to increase the sensitivity for detection of abnormal binocular saccadic eye movements, particularly of the internuclear ophthalmoplegia (INO) type associated with multiple sclerosis (MS). Saccades of 10 and 20 degrees were binocularly recorded with scleral sensor coils in 10 normal control subjects and 26 patients with definite or probable MS, seven of whom had a clinically manifest INO in one or both directions. In the cases in which this was accompanied by a dissociated nystagmus of the abducting eye, our recordings showed that such secondary saccades were also expressed, in a strongly reduced form, by the adducting eye. The patients with manifest INO showed lower average peak velocities and peak accelerations, especially for adduction of the eye on the affected side, but the distribution of these parameters overlapped with the normal distribution. A much sharper distinction between normals and patients with INOs was found by considering the ratios between peak accelerations and velocities of saccade pairs (abducting eye/adducting eye). These ratios, which eliminate much intra- and inter-individual variability, had a narrow range in normals, and all values for INOs were outside this range. On this basis, the 19 patients without clinically manifest INO were easily separated into subgroups of 14 patients with completely normal interocular ratios and five patients with elevated peak velocity and acceleration ratios, identified as sub-clinical (uni- or bilateral) INOs. Measurements of vertical saccades and of interocular timing differences provided no useful criteria for disturbances of binocular coordination in MS. We conclude that in particular, the acceleration of the adducting eye is strongly reduced in patients with an INO, and that this reduction is best identified by interocular comparison between binocular pairs of saccades.

Abstract: Vestibulo-ocular reflexes (VOR) were evaluated with a reactive torque helmet that imposed high-frequency oscillation (2-20 Hz) or step displacements of the head in the horizontal plane. The present paper describes the experimental and analytical techniques and the results for normal subjects, which will serve as a baseline for the evaluation of vestibular pathology. For comparison, manually controlled head steps were also applied, as described in the literature. Eye and head movements were recorded with magnetic search coils. Non-vestibular effects were avoided by the use of high stimulus frequencies and the analysis of short time-windows (< 100 msec) after steps. Helmet-imposed steps caused a virtually uniform head acceleration (average magnitude 770 degrees/sec) in the first 90 msec. This resulted in a linear relation between eye and head velocities; the gain and delay of the VOR could be calculated independently from the slope and offset of this relation. Such estimates appear more reliable than those obtained with conventional techniques. Normal subjects had a VOR gain of about 0.9 and a delay of about 5 msec. The results of sinusoidal head oscillation were in agreement with the results for steps. The responses to manually generated head steps agreed in general with those to helmet-induced steps, but because of the non-uniform acceleration they allowed a less exact analysis of function.

Abstract: Vestibulo-ocular reflexes (VOR) were evaluated with a reactive torque helmet that imposed high-frequency oscillation (2-20 Hz) or step displacements of the head in the horizontal plane. The present paper describes the results in patients with vestibular deficiencies (labyrinthine defective; LD); experimental and analytical techniques and results for normal subjects were described in Part 1 of this paper. The patient groups included: total unilateral LD (related to acoustic neuroma; n = 40); severe (clinically total) bilateral LD (n = 7); bilateral hyporeflexia (n = 14); unilateral hyporeflexia (n = 11); and patients with LD phenomena that had subsided (n = 3). Helmet-induced head steps provided the most specific information. Characteristically, gain was lowered in one direction or both directions after unilateral or bilateral vestibular lesions, respectively; in general, the magnitude of the gain reduction correlated well with the degree of complaints and disability. Surprisingly, delay was systematically prolonged (up to several tens of milliseconds) in all groups of subjects with manifest vestibular pathology. These results suggest that the determination of delay, in addition to gain of the VOR, is feasible and important in the evaluation of vestibular function. The results of head oscillation generally supported the results for steps, but were somewhat less specific. The responses to manually generated head steps roughly agreed with those to helmet-induced steps, but because of the non-uniform acceleration they allowed a less exact analysis of VOR function.

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Abstract: This study addresses a possible mechanism for fast disconjugate adaptation of binocular horizontal saccades. Disconjugacy of binocular saccades was elicited by two dichoptically presented, identical but aniseikonic, random checkerboard patterns. Adaptation was achieved with the patterns at far distance (144 cm). In this condition, which requires a relatively small (8%) size difference of the saccades, a short learning period was mandatory for the binocular saccades to become disconjugate. The saccadic modifications were superimposed on an idiosyncratic pattern of intra-saccadic yoking. A model of saccadic signal generation is described, that has been used to separate the contributions on saccadic disconjugacy provided by modification of visual inputs processing, which alters the motor-system inputs, and by modification of the control system: the adaptation. We identified three major components of the saccadic command (two phasic and one tonic) that contribute and in a specific way to the saccadic yoking and disconjugacy. The model analysis proposes that separate control mechanisms exist operating on these phasic and tonic signals. We show that the saccadic system can generate the vergence component shown by our aniseikonic saccades. We discuss a distributed-parallel implementation of the saccadic system able to provide both the conjugate and disconjugate components of control.

Abstract: This study describes differences in horizontal and vertical disconjugate saccades under far and close viewing conditions of two dichoptically presented aniseikonic random checkerboard patterns. At far viewing, disconjugacy of horizontal saccades requiring divergence was accomplished intrasaccadically after several minutes; for convergence the intrasaccadic disconjugacy was limited. Size differences partially persisted in open-loop trials. At close viewing intrasaccadic divergent changes in conjugacy were instantaneous, but motor storage during open-loop was absent. It is concluded that disconjugate saccades to targets at far distance lead to an adaptation process, whereas at close viewing distance horizontal disparity is a visual compensation process used directly to scale the relative amplitudes of both eyes, not leading to adaptation. The time-course of disconjugate vertical saccades was much slower, with mostly postsaccadic vergence. Nearby viewing enhanced the disconjugacy of vertical saccades.

Abstract: We investigated spontaneous variation of binocular torsion. Variation was expressed as SD of torsional eye positions measured over periods up to 32 sec. Subjects viewed a single dot target for periods of 32 sec. In half of the trials a large random-dot background pattern was superimposed on the dot. The movements of both eyes were measured with scleral induction coils. Spontaneous torsional movements were largely conjugate: cyclovergence was much more stable than cycloversion. This difference was not due to roll head movements. Stability of cyclovergence was improved by the background pattern. Although overall stability (SD of position) of cycloversion was unaffected by a background, the background induced or enhanced a small-amplitude torsional nystagmus in 3 out of 4 subjects. We hypothesize that the difference in stability of cycloversion vs cyclovergence reflects the greater importance of torsional retinal correspondence, compared to absolute torsional position. In two subjects we found evidence for the existence of cyclophoria, manifested by systematic shifts in cyclovergence caused by the appearance and disappearance of the background.

Abstract: We studied binocular cyclorotatory (torsional) eye movements in response to gratings that oscillated sinusoidally in a frontal plane. The square-wave gratings viewed by the right and left eye were presented and controlled separately to induce cycloversion and cyclovergence by oscillation in phase and out of phase. Eye movements were recorded with scleral induction coils. Stimulus oscillation frequency ranged from 0.125 to 1 Hz and the wavelength of the gratings ranged from 0.92 to 25.75 deg of visual angle. Cycloversion and cyclovergence gain were, on average, comparable in magnitude and decreased with increasing oscillation frequency. There was no consistent effect of the wavelength on the magnitude of the responses. In general, responses were considerably higher to gratings that were oriented horizontally than to those oriented vertically. This anisotropy was present both in cycloversion and cyclovergence. It was enhanced in a larger sized stimulus and by presenting stationary, orthogonal contours (mimicking a "shear" movement), but it was not consistently influenced by wavelength. Cyclovergence showed a phase lag, which increased with oscillation frequency but which was independent of wavelength. In contrast, cycloversion showed a slight phase lead which was independent of both oscillation frequency and wavelength.

Abstract: 1. The three-dimensional, binocular eye movements evoked by electrical microstimulation of the cerebellar flocculus of alert, pigmented rabbits were recorded using the scleral search coil technique. The components of these eye movements were obtained in reference to an orthogonal coordinate system consisting of a vertical axis and two horizontal axes at 45 degrees and 135 degrees azimuth. The azimuth coordinate was taken to increase to both sides from the 0 degrees reference in the direction of the nose. 2. Eye movements were evoked most readily by stimulation (0.2-ms pulses at 200 Hz for 1 s, intensity < or = 20 microA) at loci in the deep granular layer and the white matter. They consisted of slow (5-20 deg/s) movements. The responses were either binocular, with the eye ipsilateral to the stimulated flocculus usually having the larger amplitude, or were monocular, in which case they were restricted to the ipsilateral eye. 3. The evoked responses were classified according to the combination of the largest measured component of rotation for the two eyes and its sense of rotation (clockwise, CW, or counterclockwise, CCW). Seventy-eight percent of the evoked eye movements could be placed in one of two classes. For one of these classes the largest response component was a short-latency abduction of the ipsilateral eye about its vertical axis (19%), whereas for the other class (59%), the largest response component was a short-latency CCW rotation of the ipsilateral (left) eye about its 135 degrees axis. This response was frequently (50%) accompanied by a smaller short-latency CW rotation of the contralateral (right) eye about its 45 degrees axis. 4. The two main classes of three-dimensional eye movements are associated differentially with anatomically distinguishable compartments that are revealed by acetylcholinesterase histochemistry. Of the five anatomically distinguishable compartments in the floccular white matter, three are predominant. The middle of these three compartments is associated with the vertical axis class of movements, whereas the two adjacent compartments are associated with the 135 degrees class of eye movements. The eye movement relation of the other two, smaller compartments, was not determined. 5. The spatial orientation of the rotation axes of the two main classes of evoked eye movements closely corresponds to that of the preferred axes of the visual climbing fiber input to the flocculus. This suggests that both are organized in a similar coordinate system.(ABSTRACT TRUNCATED AT 400 WORDS)

Abstract: In previous work, we have demonstrated an acceleration of the buildup of slow-phase velocity of optokinetic nystagmus (OKN) after bilateral floccular injection of the aselective cholinergic agonist carbachol (Tan and Collewijn 1991; Tan et al. 1992a). In the present study we investigated the effects of unilateral floccular injections of carbachol. Such unilateral injections specifically enhanced the buildup of OKN slow-phase velocity in the direction toward the injected flocculus (ipsiversive). During binocular optokinetic stimulation, this enhancement was expressed in the motion of both eyes. Acceleration of the eye contralateral to the injected flocculus increased from 1 to about 2 degrees/s2, while the acceleration of the ipsilateral eye increased from 1 to about 1.5 degrees/s2. In contrast, buildup of contraversive OKN was unchanged. No changes were found in the steady-state OKN and optokinetic afternystagmus (OKAN). Monocular optokinetic stimulation was only effective in the nasal direction, and the effects of unilateral injection of carbachol were disconjugate. Ipsiversive OKN was enhanced only in the contralateral, seeing eye, while the response of the ipsilateral, covered eye was unchanged. We hypothesize that the directionally specific effect of unilateral cholinergic floccular stimulation on OKN is due to enhancement of predominantly the excitatory phase of modulation of the Purkinje cell's simple-spike activity by carbachol, without a marked effect of carbachol on the inhibitory phase of simple-spike modulation.

Abstract: 1. Three-dimensional rotations of both eyes were measured in alert rabbits during optokinetic stimulation about axes lying in the horizontal plane or about an earth-vertical axis, with either one or both eyes viewing the stimulus. Optokinetic stimulus speed was 2 degrees /s, either continuous or alternating in polarity (triangular stimulus). In addition to the gains of the responses, the orientations of the response axes relative to the stimulus axes were determined. 2. In comparison to the response to constant-speed optokinetic stimulation about the vertical axis, the response to constant-speed optokinetic stimulation about horizontal axes was characterized by the lack of a speed buildup. In many cases, slow phase tracking was good as long as the eye was within the central oculomotor range but deteriorated when eye deviation became more eccentric and fast phases failed to be generated. These features suggest that the optokinetic reflex about horizontal axes functions as a position-control system, rather than as a velocity-control system. 3. Binocular optokinetic stimulation at constant speed (2 degrees/s) about the roll axis (0 degrees azimuth horizontal axis) elicited disconjugate responses. Although the gain of the response was not significantly different in the two eyes (0.38 for downward and 0.44 for upward stimulation), the response axes of the two eyes differed by as much as 51 degrees. 4. Monocular, horizontal axis optokinetic stimulation at constant speed elicited responses that were grossly dissociated between the two eyes. The magnitude of the responses was anisotropic in that it varied with the azimuthal orientation of the stimulus axis; the maximum gain for each eye (0.41 for the seeing and 0.33 for the covered eye) was at 135 degrees azimuth for each eye. The axis orientation and direction (sense of rotation) of the optokinetic stimulus eliciting the maximal response for each eye coincided with the optic flow normally associated with the maximal excitation of the corresponding ipsilateral anterior canal. 5. Binocular, triangular optokinetic stimulation with small excursions (+/- 10 degrees), which avoided the saturation problems of constant-speed stimulation, elicited adequate responses without systematic directional asymmetries. Gain was approximately 0.9 for all stimulus axis orientations in the horizontal plane. 6. During monocular stimulation with triangular stimuli, the response of the seeing eye showed a gain of approximately 0.5 for all orientations of the stimulus axis. In contrast, the covered eye showed anisotropic responses, with a maximum gain of approximately 0.5 during stimulation of the seeing eye about its 45 degree axis.(ABSTRACT TRUNCATED AT 400 WORDS)

Abstract: It has been proposed that a common velocity-storage mechanism is responsible for the prolongation of vestibular nystagmus beyond the duration of the change in firing frequency of primary vestibular fibers in response to a step in velocity, and for the production of optokinetic afternystagmus (OKAN). In a previous study, bilateral injection of the aselective cholinergic agonist carbachol in the flocculus shortened the duration of buildup of optokinetic nystagmus (OKN) and the duration of OKAN, suggesting floccular involvement in velocity storage (Tan et al. 1992). In extension to that study of OKN, the present study assesses the effects of floccular carbachol on vestibular nystagmus in response to velocity steps. Our results show that injection of carbachol shortens the duration of vestibular nystagmus from about 13 to about 8 s; a finding which supports a common velocity-storage mechanism for optokinetic and vestibular signals. We propose that the indistinguishable effects of carbachol on OKAN and vestibular nystagmus are due to modification of the transmission of an oculomotor corollary signal, which has been identified electrophysiologically in the flocculus.

Abstract: Congenital nystagmus (CN) has been described as a 'fixation' nystagmus implying an inability to fixate a target. However, each cycle of CN contains a target-foveation period during which the eye velocity is at, or near, zero. Prolongation of foveation time, reduction of retinal image velocity and cycle-to-cycle foveation repeatability all contribute to increased visual acuity. We developed several methods to accurately measure the dynamics of foveation in CN; their use is illustrated on an individual with typical idiopathic CN and no afferent defects. During eight 5-second intervals of fixation on a stationary target, the horizontal standard deviation (SD) of the mean foveation position (FPOS) was 12.82 minarc and the SD of foveation velocity was 118.36 minarc/sec. The SD of the means of total eye position and of the non-foveating peak of the CN were 43.17 and 25.32 minarc respectively. The mean foveation-time interval (eye velocity less than or equal to 4 degrees/sec) was 57.27 msec. The SD FPOS for the best 1-second interval (4 successive CN cycles), in a typical 5-second record, was 0.71 minarc. Histograms revealed peaks of eye position at 0 +/- 10 minarc and of eye velocity at 0 +/- 240 minarc/sec. The small vertical component of the CN (16 minarc peak-to-peak) had a SD of 6.56 minarc. A nystagmus foveation function related to visual acuity was derived that was more sensitive than CN intensity. The increased visual acuity resulting from the use of convergence or base-out prisms was due to increased foveation time. Although it might appear that CN is a defect of fixation, this individual with CN had strong fixation reflexes in the sense that he was able to accurately (within 1 minarc) achieve (interbeat) and maintain (intrabeat) target foveation for appreciable periods of time. Our data support the hypothesis that individuals with idiopathic CN do not have a primary disturbance of fixation.

Abstract: It has been shown that, during 5 seconds of fixation, an individual with congenital nystagmus (CN) can repeatedly (beat-to-beat) foveate (SD = 12.87 minarc) and maintain low retinal slip velocities (SD = 118.36 minarc/sec). Smooth pursuit data from several CN subjects showed that eye velocities during these foveation intervals approximated target velocity. Despite some claims that CN is caused by absent or "reversed" smooth pursuit, those with CN hardly ever experience oscillopsia or exhibit any accompanying symptoms of such deficits in pursuit; they are able to master sports requiring tracking of rapidly moving small objects (e.g. racquetball or handball). We developed and describe several new methods to accurately assess the function of smooth pursuit in an individual with typical idiopathic CN. We investigated the dynamics of CN foveation periods during smooth pursuit to test the hypothesis that eye velocities would match target velocities during these periods. Unity or near-unity instantaneous (beat-to-beat) pursuit gains of both experimenter-moved and subject-moved targets at peak velocities ranging from only a few deg/sec up to 210 degrees/sec were measured. The dynamic neutral zone was found to shift oppositely to target direction by amounts proportional to the increase in target speed. Our methods proved that eye velocity is made to match target velocity during the foveation intervals and support the conclusion that smooth pursuit in individuals with CN is functioning normally in the presence of the CN oscillation. In addition, we hypothesize that the same fixation mechanism that prevents oscillopsia during fixation of stationary targets, also does so during pursuit.

Abstract: 1. In the alert, pigmented rabbit, eye movements were recorded during optokinetic nystagmus (OKN) and during optokinetic after nystagmus (OKAN). These responses were elicited by steps in surround-velocity ranging from 5-110 degrees/s during binocular as well as monocular viewing. 2. In the baseline condition, OKN showed an approximately linear build-up of eye velocity to a steady-state, followed by a linear decay of eye velocity during OKAN after the lights were turned off. Build-up during binocular viewing was characterized by a constant, maximum eye-acceleration (about 1 degree/s2) for stimulus velocities up to 60 degrees/s. OKAN, instead, was characterized by a fixed duration (about 10 s) for stimulus velocities up to 20 degrees/s. Steady-state eye velocity saturated at about 50 degrees/s. 3. Monocular stimulation in the preferred (nasal) direction elicited a build-up that was on average twice as slow as during binocular stimulation. Steady-state velocity during monocular stimulation saturated at about 20 degrees/s. OKAN was of equal duration as during binocular stimulation. In the non-preferred direction, a very irregular nystagmus was elicited without velocity build-up. The stronger response to binocular stimulation, compared to the responses under monocular viewing condition in either nasal and temporal direction suggests potentiation of the signals of either eye during binocular viewing. 4. OKN and OKAN were re-assessed after intra-floccular micro-injection of the nonselective cholinergic agonist carbachol. In the binocular viewing condition, eye-acceleration during build-up was strongly enhanced from 1 degree/s2 before to 2.5 degrees/s2 after injection. The saturation level of steady-state eye velocity was also increased, from 50 degrees/s before to more than 60 degrees/s after carbachol. The duration of OKAN, however, was shortened from 10 s before to 6 s after injection. The response to monocular stimulation in the preferred direction revealed similar changes. 5. The flocculus appears to be involved in the control of the dynamics of OKN in the rabbit. Cholinergic mechanisms affect the floccular control of the rate at which slow-phase velocity can be built up and the rate of decay of eye velocity during OKAN. Cholinergic stimulation of the flocculus enhances the dynamics of OKN, while velocity storage is shortened.

Abstract: Bilateral microinjections into the cerebellar flocculus of the rabbit of carbachol, a general cholinergic agonist, profoundly affect vestibuloocular (VOR) and optokinetic (OKR) reflexes. For sinusoidal stimuli (0.15 Hz, 5 deg peak to peak), the gain of the OKR was strongly increased, while the gain of the VOR was moderately increased. These effects were partially mimicked by floccular injection of the acetylcholinesterase inhibitor eserine. Floccular injection of the muscarinic blocker atropine significantly lowered the gain of the OKR. The effects of the nicotinic blocker mecamylamine were not significant. Optokinetic nystagmus (OKN) in response to constant stimulus velocities (1-30 deg/second) showed a markedly accelerated buildup and a shortened optokinetic after-nystagmus (OKAN) after floccular injections of carbachol. The steady-state gain of OKN remained unaffected. None of the described effects occurred after floccular injection of the solvent, saline. It is postulated that cholinergic cerebellar afferents, one probable source of which are the vestibular nuclei, enhance the optokinetic and vestibular modulation of floccular Purkinje cells.

Abstract: Binocular cyclorotatory (torsional) eye movements in response to visual patterns, which oscillated sinusoidally in the frontal plane, were recorded with scleral induction coils in human subjects. Conjugate cycloversion and disjunctive cyclovergence were directly compared by in-phase and out-of-phase oscillation of the same pattern. Stimulus motion had a frequency of 0.2 Hz and amplitudes of 2-8 deg. Both response types had a similar and low gain (about 0.2 averaged over all subjects). Cycloversion showed no time lag, while cyclovergence lagged by about 600 msec. Non-fusible patterns were effective in eliciting cycloversion, but not cyclovergence. Apart from this, the nature of the pattern (randomly distributed dots, regular rows of dots, horizontal or vertical grating, Julesz stereogram or images with a pictorial significance) had only the slightest effect on the magnitude of the responses.

Abstract: It has been shown that, during fixation of a stationary target with a fixed head, an individual with congenital nystagmus (CN) can repeatedly (beat-to-beat) foveate (within 13 minarc) and maintain low retinal slip velocities (less than 4 degrees/sec). With the head in motion, vestibuloocular reflex (VOR) data showed eye velocities during these foveation periods that approximation head velocity. Despite some claims that the VOR of CN subjects was deficient or absent, individuals with CN hardly ever complain of oscillopsia or exhibit any of the symptoms that would accompany such deficits in the VOR, whether during simple walking and running or while skiing down a mogul field. We developed and describe several different and unrelated methods to accurately assess the function of the VOR in an individual with typical idiopathic CN. We investigated the dynamics of CN foveation periods during head rotation to test the hypothesis that eye velocities would match head velocities during these periods. At about 1 Hz, horizontal VOR instantaneous (beat-to-beat) gains were 0.96 in the light and 0.94 in the dark while imaging a stationary target. Vertical VOR gains were 1.00 and 0.99 for these two conditions at the same frequency; the CN was horizontal. Also, during the VOR there is a CN neutral-zone shift comparable to that found during smooth pursuit. Our methods demonstrated that gaze velocity was held constant during foveation periods and we conclude that the VOR in this subject is functioning normally in the presence of the CN oscillation. Based on our findings in this and previous studies, we hypothesize that CN may be due to a peripheral instability.

Abstract:

Abstract: Noradrenaline (NA) has been implicated as a neuromodulator in plasticity, presumably facilitating adaptive processes. Since the flocculus receives noradrenergic afferents, and ablation of the flocculus interferes with the normal adaptive changes in the VOR gain, experiments were performed to find out whether bilateral injection of monoaminergic substances into the flocculus of rabbits could modify the adaptive changes of the VOR. The visual world surrounding the rabbit was oscillated in opposite direction to the platform on which the rabbit was mounted, which resulted in an adaptive increase in the VOR gain; this adaptation was measured either in light or in darkness. Floccular injection of the beta-agonist isoproterenol did not greatly affect the adaptation of the VOR measured in light. In darkness, however, the increase in gain after injection of isoproterenol was larger than during normal adaptation. The beta-antagonist sotalol reduced the adaptation of the VOR gain significantly in light as well as in darkness. In a control condition without pressure for adaptation (only intermittent testing of the VOR gain over a period of 2.5 h), the gain of the VOR was not significantly affected by similar injections of beta-adrenergic agents. We conclude that the noradrenergic system facilitates the adaptation of the VOR gain to retinal slip in rabbits without affecting the VOR gain directly. At least part of this influence is exerted through beta-receptors located in the cerebellar flocculus.

Abstract: Noradrenaline (NA) has been implicated as a neuromodulator in plasticity, presumably facilitating adaptive processes. Recent experiments by others have suggested a modulatory role of NA in adaptive changes in the vestibulo-ocular reflex (VOR). These experiments showed that general depletion of brain NA resulted in a decreased ability to produce adaptive changes in the VOR gain. In order to identify the specific brain region responsible for these effects, as well as the nature of the adrenoceptors involved, we injected beta-adrenergic substances bilaterally into the flocculus of rabbits. The flocculus is known to receive noradrenergic afferents and, moreover, ablation of the flocculus interferes strongly with the normal adaptive changes in the VOR gain. We injected the beta-agonist isoproterenol and the beta-antagonist sotalol, and compared the adaptive capacity of the rabbits after these injections to that in a situation without injection. The rabbit was oscillated in a direction opposite to the direction of motion of the platform on which the rabbit was mounted, a condition which normally results in an increase in the VOR gain, measured either in light or in darkness. Injection of the beta-agonist did not greatly affect the adaptation of the VOR measured in the light. In darkness, the increase in gain after the injection of isoproterenol was larger than in the non-injection experiments in 9 out of 10 rabbits. The beta-antagonist sotalol reduced the adaptation of the VOR gain significantly in the light, as well as in darkness. In a control condition without pressure for adaptation (only intermittent testing of the VOR gain over a period of 2.5 h), the gain of the VOR either remained unaffected or was only slightly affected by similar injections of beta-adrenergic agents in individual rabbits. For the group as a whole, these effects were insignificant. We conclude from these results that noradrenergic systems facilitate the adaptation of the VOR gain to retinal slip in rabbits, without affecting the VOR gain directly. At least part of this influence is exerted through beta-receptors located in the cerebellar flocculus.

Abstract: Rabbits trained to discriminate vertical vs. oblique striations are unable to discriminate angular differences of 5 degrees. In the present study the instabilities around the roll axis of the eyes were measured during visual discrimination. The results indicate that these instabilities are one of the causes of the rabbit's inability to discriminate angular differences of 5 degrees.

Abstract: The climbing fibres (CFs) of the rabbit flocculus that respond in a speed- and direction-selective manner to retinal image slip produced by eye rotations can be divided into three classes on the basis of the orientation of the rotation axis associated with their greatest modulation (the preferred axis). The similarity of the orientations of these axes to those of the eye rotation axes of the extraocular muscles suggests that a simple geometrical correspondence may exist between the eye rotation associated with the preferred axis of a given class of CFs and the eye rotation produced by activation of the Purkinje cells upon which that class of CFs synapse. To pursue this possibility, the axes of the eye rotations evoked by electrical microstimulation of the alert rabbit's flocculus were determined simultaneously for both eyes in three dimensions using two orthogonal search coils on each eye. A limited number of slow eye movement response patterns were found, and of these, two predominated. The most common response was a counterclockwise (CCW) rotation of the ipsilateral (left) eye around an axis close to the horizontal plane and at about 140 degrees posterior to the nose. The other predominant response was abduction of the ipsilateral eye. These two response patterns, together with the smaller conjugate components for the contralateral eye, are consonant with the orientations of the preferred CF axes. In addition, a clear CCW rotation of the contralateral (right) eye about its 135 degrees axis was also evoked from some stimulation sites. This response, which occurred either alone or as a component of an upward rotation about the nasal-occipital (roll) axis, is at variance with the orientations of the preferred CF axis. However, the latencies of the CCW contralateral 135 degrees component (80-140 ms) were greater than those of the CW contralateral 45 degrees component, the CCW ipsilateral 135 degrees component and the ipsilateral abduction component (8-48 ms). These latency differences may distinguish stimulation of Purkinje cells from stimulation of other neurones.

Abstract: Horizontal binocular eye movements of four subjects were recorded with the scleral sensor coil--revolving magnetic field technique while they fixated a natural target, whose distance was varied in a normally illuminated room. The distance of the target relative to the head of the subject was changed in three ways: (a) the target was moved manually by the experimenter; (b) the target was moved manually by the subject; (c) the target remained stationary while the subject moved his upper torso towards and away from the target. The rate of change of target distance was varied systematically in four levels, ranging from 'slow' to 'very fast', corresponding to changes in target vergence from about 10 degrees s-1 to about 100 degrees s-1. The dynamics of ocular vergence with regard to delay and speed were, under all three conditions, considerably better than could be expected from the literature on ocular vergence induced by disparity and/or blur. When 'very fast' changes in the distance of the target were made, subjects achieved maximum vergence speeds of up to about 100 degrees s-1. Delays of these fast vergence responses were generally smaller than 125 ms. Negative delays, i.e. ocular vergence leading the change in target distance, were observed. The eyes led the target (i.e. predicted target motion) by about 90 ms on average, when the subject used his hand to move the target. Vergence tracking was almost perfect when changes in distance were produced by moving the upper torso. In this condition, the eye led the target by about 5 ms. In the 'slow' and 'medium' conditions (stimulus speeds about 10-40 degrees s-1) tracking was accurate to within 1-2 degrees, irrespective of the way in which the target was moved. In the 'fast' and 'very fast' conditions (stimulus speeds about 40-100 degrees s-1), the accuracy of vergence tracking was better for self-induced than for experimenter-induced target displacements, and accuracy was best during voluntary movements of the upper torso. In the last case, ocular vergence speed was within about 10% of the rate of change of the vergence angle formed by the eyes and the stationary target. The dynamics of convergent and divergent vergence responses varied considerably. These variations were idiosyncratic. They were consistent within, but not between, subjects. Ocular vergence associated with attempted fixation of an imagined target, changing distance in darkness, could only be made by two of the four subjects.(ABSTRACT TRUNCATED AT 400 WORDS)

Abstract: Anatomical and physiological findings indicate that the crossed optic fibres of the rabbit have a crucial role in binocular vision. In order to directly examine the visual functions of the uncrossed fibre system, a technique of sectioning the optic chiasma midsagitally was developed. Both normal and chiasma-sectioned rabbits were tested on a variety of visual discrimination tasks as well as such oculomotor control functions as the optokinetic and vestibulo-ocular reflexes. Following transection of all contralateral retinal projections, rabbits were found to retain the same visual capacity for detection of intensity and orientation differences as before the operation. There was, however, a complete loss of optokinetic reflexes and a 50% reduction of the vestibulo-ocular reflex both in the light and in the dark.

Abstract: We studied the effects of unilateral frontal eye-field (FEF) lesions on eye-head coordination in monkeys that were trained to perform a visual search task. Eye and head movements were recorded with the scleral search coil technique using phase angle detection in a homogeneous electromagnetic field. In the visual search task all three animals showed a neglect for stimuli presented in the field contralateral to the lesion. In two animals the neglect disappeared within 2-3 wk. One animal had a lasting deficit. We found that FEF lesions that are restricted to area 8 cause only temporary deficits in eye and head movements. Up to a week after the lesion the animals had a strong preference to direct gaze and head to the side ipsilateral to the lesion. Animals tracked objects in contralateral space with combined eye and head movements, but failed to do this with the eyes alone. It was found that within a few days after the lesion, eye and head movements in the direction of the target were initiated, but they were inadequate and had long latencies. Within 1 wk latencies had regained preoperative values. Parallel with the recovery on the behavioral task, head movements became more prominent than before the lesion. Four weeks after the lesion, peak velocity of the head movement had increased by a factor of two, whereas the duration showed a twofold decrease compared with head movements before the lesion. No effects were seen on the duration and peak velocity of gaze. After the recovery on the behavioral task had stabilized, a relative neglect in the hemifield contralateral to the lesion could still be demonstrated by simultaneously presenting two stimuli in the left and right visual hemifields. The neglect is not due to a sensory deficit, but to a disorder of programming. The recovery from unilateral neglect after a FEF lesion is the result of a different orienting behavior, in which head movements become more important. It is concluded that the FEF plays an important role in the organization and coordination of eye and head movements and that lesions of this area result in subtle but permanent changes in eye-head coordination.

Abstract: Previously we reported that failures of compensatory eye movements led to appreciable binocular retinal image motion during head rotation. Subjectively, the visual world appeared clear, fused, and stable under these conditions. The present experiments examined these impressions psychophysically. The spatial modulation transfer function of subjects with known retinal image motion was measured during head rotation. We found that contrast sensitivity was reduced for gratings over 6 cycles/degree and was increased for lower spatial frequencies. Our results, when compared with Kelly's [J. Opt. Soc. Am. 69, 1340-1349 (1979)] measurements made with artificially moving stabilized gratings, show that natural retinal image motion is less harmful to contrast sensitivity at high spatial frequencies and more beneficial at low spatial frequencies. Furthermore, we had previously found that natural retinal image motion was different in each eye during head movement but no diplopia was noticed. We confirmed this subjective impression by measuring forced-choice stereoacuity thresholds concurrent with binocular head and eye recordings. Stereoacuity was not disturbed by large fixation disparities or high vergence velocities. Recordings also were made while a fused Julesz stereogram was viewed during attempts to break fusion with violent head movements. Fusion could not be broken. Stereograms turned on during violent head movement fused rapidly. We conclude that vision is better with natural retinal image motion than expected from experiments done with stabilized heads.

Abstract: Static and dynamic components of ocular counterroll as well as cyclorotatory optokinetic nystagmus were measured with a scleral search coil technique. Static counterroll compensated for about 10% of head roll when the head was tilted to steady positions up to 20 deg from the upright position. The dynamic component of counterroll, which occurs only while the head is moving, is much larger. It consists of smooth compensatory cyclorotation opposite to the head rotation, interrupted frequently by saccades moving in the same direction as the head. During voluntary sinusoidal head roll, cyclorotation compensated from 40% to more than 70% of the head motion. In the range 0.16 to 1.33 Hz, gain increased with frequency and with the amount of visual information. The lowest values were found in darkness. The gain increased in the presence of a visual fixation point and a further rise was induced by a structured visual pattern. Resetting saccades were made more frequently in the dark than in the light. These saccades were somewhat slower than typical horizontal saccades. Cyclorotatory optokinetic nystagmus could be induced by a patterned disk rotating around the visual axis. It was highly variable even within a same subject and had in general a very low gain (mean value about 0.03 for stimulus velocities up to 30 deg/s). It is concluded that cyclorotational slip velocity on the retina is considerably reduced by counterroll during roll of the head, although the residual cyclorotation after the head has reached a steady position is very small.

Abstract: Eye movements associated with eyelid closure were recorded in human subjects with search coils, embedded in self-adhering scleral annuli, in a magnetic field. In contrast to classical notions, voluntary as well as reflex blinks were consistently accompanied by transient downward and nasalward movements of both eyes with amplitudes 1-5 degrees. These eye movements had a shorter duration than the upper lid movements, and the shapes of the spatial trajectories of eye and lid movements were not similar. The trajectory of the eye movements was only modestly affected by gaze eccentricities up to 15 degrees; there was a tendency for the downward component to be enhanced by looking upward, and vice versa. Restraining of the lids of one eye in the open or closed position did not significantly alter the eye movements during (attempted) blinks. Velocity-amplitude-duration relations of the down- and upward components were similar for the same eye before and after closure and for the closed eye and the contralateral unrestrained eye. The velocity-amplitude-duration characteristics of saccades were also unaffected by prolonged closure of the lids of one eye. Prolonged, voluntary closure of the lids was followed by a slow, tonic ocular deviation, which was consistently upward in half of the subjects and consistently downward in the other half. Additional horizontal components were highly variable even within subjects. In one subject the downward deviation was converted into upward deviation when lid closure was mechanically impeded. We conclude that elevation of the eye ball (Bell's phenomenon) does not occur during short blinks and only in about half of the subjects during voluntary unrestrained prolonged lid closure. Our evidence does not support the possibility that the transient eye movements during blinks are caused primarily by a mechanical interaction between the lids and the eye (or the scleral annulus). More likely, they are a secondary effect of an active cocontraction of extraocular muscles that primarily results in retraction of the eye.

Abstract: Some neurones in macaque postarcuate premotor area modulate their firing frequency in relation to motor tasks which require visual information. We previously reported that a large proportion of these neurones modulate during execution of a detour reaching task in which the movement phase was separated in time from the phase in which the monkey received a visual cue for the movement required to retrieve a food reward. A large proportion of task-related neurones (75%) modulated during this 'visual' phase, in which no task-related movements were made. This modulation was related to the position of the food reward, which served as the visual cue. Most of these neurones were located in cortical area 6, close to the arcuate curvature and its spur, but also more caudally in area 4 and rostrally in area 8. In the present chronic recording experiments in monkeys, several variations of the original task were used in order to test whether the 'visual'-related neuronal modulation could be involved in preparation of the upcoming movement. This modulation is unlikely to be related to any eye or arm movements occurring during the visual phase or to changes in environmental illumination. Neither can it be related to the presence of the visual cue in a particular part of the visual field, since the pattern of neuronal modulation was similar when a cue with a fixed position was used. This modulation was, however, contingent upon the occurrence of food retrieval during the subsequent 'movement phase', since it was abolished or diminished during presentation of a 'food-reward' which the monkey did not retrieve. For several neurones, modulation pattern during the visual phase depended on whether the food reward was to be retrieved with a gross hand movement or with relatively independent finger movements. It is likely, therefore, that neurones in the postarcuate premotor cortex are involved in preparation for arm movements with the help of visual cues. The results are discussed in view of corticocortical pathways which might be involved in transmission of visual information from visual areas through parietal association areas and premotor cortex to the primary motor cortex.

Abstract: Prior work has shown that smooth eye movements in the presence of both stationary and moving stimuli are determined, at least in part, by the voluntary selection of either the stationary or the moving stimulus as the target for smooth eye movements. The effectiveness of voluntary selection in eliminating the influence of the stimuli not selected (i.e. backgrounds) on smooth eye movement is not known because prior studies used targets and backgrounds with different physical characteristics. Thus, effects of voluntary selection were confounded with the relative strength of target and background as stimuli for smooth eye movements. We measured eye movements (resolution 1') of two highly-experienced eye movement subjects with a target and background with the same physical characteristics: two, identical, full-field, superimposed patterns of randomly-positioned dots (1 dot/deg2 or 8 dots/deg2). One field was stationary and the other moved at 70.2 minarc/sec. The effect of the moving background on smooth eye movements when the stationary field was the target, and the effect of the stationary background on smooth eye movements when the moving field was the target was negligible (0-4% for one subject; 0-2% for the other). The influence of the background on smooth eye movements was affected by a six-fold reduction in the intensity of either the target or background, but effects of such intensity changes were small and different for each subject. Taken together, these results show that the effectiveness of voluntary selection in eliminating the influence of background stimuli on smooth eye movements can be virtually complete. Any observed influence of the background--however small--can be attributed to voluntary factors (e.g. subjects' failure to apply sufficient effort or attention) rather than to the operation of an involuntary mechanism that automatically integrates velocity information from target and background. The attention and effort required to ensure that voluntary selection is perfect may impair the accuracy of psychophysical judgments made about the background.

Abstract: Eye and head movements in the horizontal, frontal and sagittal planes were recorded in the rabbit with a newly developed technique using dual scleral search coils in a rotating magnetic field. The compensatory eye movements elicited by passive sinusoidal oscillation deteriorated for frequencies below 0.1 Hz in the horizontal, but not in the frontal and sagittal planes. In the light gain was relatively independent of frequency in all planes and amounted to 0.82-0.69, 0.92-0.83 and 0.65-0.59 in the horizontal, frontal and sagittal plane, respectively. In freely moving animals, similar input-output relations were found. The stability of the retinal image thus proved to be inversely proportional to the amount of head movements associated with behavioural activity. Maximal retinal image velocities varied between 2-4 degree/s for a rabbit sitting quietly and 30-40 degrees/s during locomotor activity. Gaze displacements showed different characteristics in the various planes, possibly in relation with the structure of the retinal visual streak. Horizontal gaze changes were mainly effected by saccades. Gaze changes in the frontal plane were relatively rare and effected by non-saccadic, combined head and eye movements with temporary suppression of compensatory eye movements. Eye rotations in the sagittal plane, possibly functioning to adjust the direction of binocular vision vertically, were abundant and effected by large head movements in combination with a low gain of compensatory eye movements in this plane.

Abstract: Pursuit of a point target in real or apparent motion upon a dark, diffusely lighted or structured background was recorded with a scleral coil technique. Smooth and saccadic components were separated and analyzed with computer techniques. Sigma-pursuit was superior to pursuit of beta- or real motion: smooth pursuit gain was higher, saccadic rate was lower and the detrimental effect of a structured background was smaller. Due to directional errors, smooth pursuit velocity often exceeded target velocity when this was smaller than about 10 degrees/sec. However, the smooth component in the correct direction of the target motion had a gain less than or equal to 1.0 and decreasing at higher target velocities for all pursuit modes, inclusive sigma-pursuit.

Abstract: The decay of the slow phase velocity of post-rotatory (PRN) and optokinetic (OKAN) after-nystagmus as a function of time was measured in Dutch rabbits after stimulation with velocity steps of 30, 60, and 150 degrees/s. The decays fitted linear functions very well, but only poorly exponential ones. Typical decay rates were 2-5 degrees/s2, with apparent time constants (defined by decay to 37% of initial velocity) in the order of 10-20 s. Within one animal, the decays of OKAN and PRN with similar initial velocities were indistinguishable. With sinusoidal oscillation, the time constant of the vestibulo-ocular reflex - estimated from phase lead - was only 2-3 s, and probably similar to the cupular time constant. In general, time constants increased when eye velocities increased. This indicates that the vestibulo-ocular reflex of the rabbit behaves as a non-linear system. A velocity storage system with a constant discharge rate is postulated as a main non-linear element. This would introduce a linear decay of velocity as well as a threshold for velocity. This storage system would be common to both vestibulo-ocular and optokinetic reflexes.