Retinal Image Position Research Articles

The effect of retinal image error update rate on human vestibulo-ocular reflex gain adaptation.

The primary function of the angular vestibulo-ocular reflex (VOR) is to stabilise images on the retina during head movements. Retinal image movement is the likely feedback signal that drives VOR modification/adaptation for different viewing contexts. However, it is not clear whether a retinal image position or velocity error is used primarily as the feedback signal. Recent studies examining this signal are limited because they used near viewing to modify the VOR. However, it is not known whether near viewing drives VOR adaptation or is a pre-programmed contextual cue that modifies the VOR. Our study is based on analysis of the VOR evoked by horizontal head impulses during an established adaptation task. Fourteen human subjects underwent incremental unilateral VOR adaptation training and were tested using the scleral search coil technique over three separate sessions. The update rate of the laser target position (source of the retinal image error signal) used to drive VOR adaptation was different for each session [50 (once every 20 ms), 20 and 15/35 Hz]. Our results show unilateral VOR adaptation occurred at 50 and 20 Hz for both the active (23.0 ± 9.6 and 11.9 ± 9.1% increase on adapting side, respectively) and passive VOR (13.5 ± 14.9, 10.4 ± 12.2%). At 15 Hz, unilateral adaptation no longer occurred in the subject group for both the active and passive VOR, whereas individually, 4/9 subjects tested at 15 Hz had significant adaptation. Our findings suggest that 1-2 retinal image position error signals every 100 ms (i.e. target position update rate 15-20 Hz) are sufficient to drive VOR adaptation.

Visual contribution to the high-frequency human angular vestibulo-ocular reflex

The vestibulo-ocular reflex (VOR) acts to maintain images stable on the retina by rotating the eyes in exactly the opposite direction, but with equal magnitude, to head velocity. When viewing a near target, this reflex has an increased response to compensate for the translation of the eyes relative to the target that acts to reduce retinal image slip. Previous studies have shown that retinal velocity error provides an important visual feedback signal to increase the low-frequency (<1 Hz) VOR response during near viewing. We sought to determine whether initial eye position and retinal image position error could provide enough information to substantially increase the high-frequency VOR gain (eye velocity/head velocity) during near viewing. Ten human subjects were tested using the scleral search coil technique during horizontal head impulses under different lighting conditions (constant dark, strobe light at 0.5, 1, 2, 4, 10, 15 Hz, constant light) while viewing near (9.5 ± 1.3 cm) and far (104 cm) targets. Our results showed that the VOR gain increased during near viewing compared to far viewing, even during constant dark. For the near target, there was an increase in VOR gain with increasing strobe frequency from 1.17 ± 0.17 in constant dark to 1.36 ± 0.27 in constant light, a 21 ± 9 % increase. For the far target, strobe frequency had no effect. Presentation order of strobe frequency (i.e. 0.5-15 vs. 15-0.5 Hz) did not affect the gain, but it did affect the vergence angle (angle between the two eye's lines of sight). The VOR gain and vergence angles were constant during each trial. Our findings show that a retinal position error signal helps increase the vergence angle and could be invoking vestibular adaptation mechanisms to increase the high-frequency VOR response during near viewing. This is in contrast to the low-frequency VOR that depends more on retinal velocity error and predictive adaptation mechanisms.

Evidence that convergence rather than accommodation controls intermittent distance exotropia

This study considered whether vergence drives accommodation or accommodation drives vergence during the control of distance exotropia for near fixation. High accommodative convergence to accommodation (AC/A) ratios are often used to explain this control, but the role of convergence to drive accommodation (the CA/C relationship) is rarely considered. Atypical CA/C characteristics could equally, or better, explain common clinical findings. Nineteen distance exotropes, aged 4-11 years, were compared while controlling their deviation with 27 non-exotropic controls aged 5-9 years. Simultaneous vergence and accommodation responses were measured to a range of targets incorporating different combinations of blur, disparity and looming cues at four fixation distances between 2 m and 33 cm. Stimulus and response AC/A and CA/C ratios were calculated. Accommodation responses for near targets (p = 0.017) and response gains (p = 0.026) were greater in the exotropes than in the controls. Despite higher clinical stimulus AC/A ratios, the distance exotropes showed lower laboratory response AC/A ratios (p = 0.02), but significantly higher CA/C ratios (p = 0.02). All the exotropes, whether the angle changed most with lenses ('controlled by accommodation') or on occlusion ('controlled by fusion'), used binocular disparity not blur as their main cue to target distance. Increased vergence demand to control intermittent distance exotropia for near also drives significantly more accommodation. Minus lens therapy is more likely to act by correcting overaccommodation driven by controlling convergence, rather than by inducing blur-driven vergence. The use of convergence as a major drive to accommodation explains many clinical characteristics of distance exotropia, including apparently high near stimulus AC/A ratios.

Perisaccadic Stereo Depth with Zero Retinal Disparity

When an object is viewed binocularly, unequal perspective projections of the two eyes' half images (binocular disparity) provide a cue for the sensation of stereo depth. For almost 200 years, binocular disparity has remained synonymous with retinal disparity, which is computed by subtracting the distance of each half image from its respective fovea. However, binocular disparity could also be coded in headcentric instead of retinal coordinates, by combining eye position and retinal image position in each eye and representing disparity as differences between visual directions of half images relative to the head. Although these two disparity-coding schemes suggest very different neural mechanisms, both offer identical predictions for stereopsis in almost every viewing condition, making it difficult to empirically distinguish between them. We designed a novel stimulus that uses perisaccadic spatial distortion to generate inconsistency between headcentric and retinal disparity. Foveal half images flashed asynchronously just before a horizontal saccade have zero retinal disparity, yet they produce a sensation of depth consistent with a nonzero headcentric disparity. Furthermore, this headcentric disparity can cancel and reverse the perceived depth stimulated with nonzero retinal disparity. This is the first demonstration that a coding scheme other than retinal disparity has a role in human stereopsis.

How color break-up occurs in the human-visual system: The mechanism of the color break-up phenomenon

— In the present set of experiments, we examined the mechanisms underlying color break-up (CBU), a phenomenon observed when images produced with a color-sequential projector are viewed. The perceived position of CBU was measured during fast eye movement, saccade with static and briefly flashed stimuli. Results showed that CBU did not simply correspond to the locus of the stimulus on the retina during saccades, because the width of the CBU perception was narrower than the distance of the eye movements. This effect was thought to be related to visual stability, which allows objects to be perceived as stationary even when the eyes move and the retinal image changes position. Visual stability is assumed to operate by compensating for the change in retinal image position using eye-position signals; however, this compensation is imperfect during saccades. Thus, incomplete compensation results in a CBU perception that is of a narrower width than the amplitude of the saccade. In conclusion, CBU cannot be simulated with moving video cameras because it results largely from the mechanisms of visual perception. Large inter-individual differences in perception of CBU were also found. This observation also supports the idea that CBU depends on the mechanism of perception.

Role of arcuate frontal cortex of monkeys in smooth pursuit eye movements. I. Basic response properties to retinal image motion and position.

Anatomical and physiological studies have shown that the "frontal pursuit area" (FPA) in the arcuate cortex of monkeys is involved in the control of smooth pursuit eye movements. To further analyze the signals carried by the FPA, we examined the activity of pursuit-related neurons recorded from a discrete region near the arcuate spur during a variety of oculomotor tasks. Pursuit neurons showed direction tuning with a wide range of preferred directions and a mean full width at half-maximum of 129 degrees. Analysis of latency using the "receiver operating characteristic" to compare responses to target motion in opposite directions showed that the directional response of 58% of FPA neurons led the initiation of pursuit, while 19% led by 25 ms or more. Analysis of neuronal responses during pursuit of a range of target velocities revealed that the sensitivity to eye velocity was larger during the initiation of pursuit than during the maintenance of pursuit, consistent with two components of firing related to image motion and eye motion. FPA neurons showed correlates of two behavioral features of pursuit documented in prior reports. 1) Eye acceleration at the initiation of pursuit declines as a function of the eccentricity of the moving target. FPA neurons show decreased firing at the initiation of pursuit in parallel with the decline in eye acceleration. This finding is consistent with prior suggestions that the FPA plays a role in modulating the gain of visual-motor transmission for pursuit. 2) A stationary eccentric cue evokes a smooth eye movement opposite in direction to the cue and enhances the pursuit evoked by subsequent target motions. Many pursuit neurons in the FPA showed weak, phasic visual responses for stationary targets and were tuned for the positions about 4 degrees eccentric on the side opposite to the preferred pursuit direction. However, few neurons (12%) responded during the preparation or execution of saccades. The responses to the stationary target could account for the behavioral effects of stationary, eccentric cues. Further analysis of the relationship between firing rate and retinal position error during pursuit in the preferred and opposite directions failed to provide evidence for a large contribution of image position to the firing of FPA neurons. We conclude that FPA processes information in terms of image and eye velocity and that it is functionally separate from the saccadic frontal eye fields, which processes information in terms of retinal image position.

Changes in fixation in the presence of prism monitored with a confocal scanning laser ophthalmoscope

Background:The main optical effect of an ophthalmic prism is to deviate uniformly the entire field of view seen through the prism, resulting in an equal eye movement, if fixation on the same object is to be maintained. Conversely, it has been suggested that, in the case of low vision due to central scotoma, the eye changes its eccentric viewing behaviour when a prism is introduced, that is, the eye remains stationary with a change of retinal image location. Method:A new method is described in which retinal image position was recorded with a confocal scanning laser ophthalmoscope in four experimental conditions. Results:Results showed that the normally‐sighted eye rotates rapidly to compensate for the introduction of a prism. Conclusion:Future studies are required involving subjects with central scotoma but it is likely that many subjects with low vision will also refixate behind a prism and that the prescribing and wearing of bilateral prism in cases of central vision loss will have limited effectiveness in the demonstration or training of eccentric viewing.

Changes in Saccadic and Manual Motor Control After Ocular Smooth Pursuit Adaptive Modifications

ABSTRACT The gain of ocular smooth pursuit responses can be adaptively modified under certain circumstances. Evidence that these modifications are caused, at least partly, by changes at a sensory level comes from the fact that subsequent manual tracking movements generated during visual fixation are also modified in a similar manner. The question addressed in the present experiment was whether the modifications are constrained specifically to responses such as these, which are driven by retinal image motion, or whether they can also influence movements that make use of retinal image position. This was accomplished by comparing manual and oculomotor responses to step changes in target position before and after smooth pursuit adaptation. The results showed that, as with manual tracking movements, these responses were also modified by the adaptive procedure, although to a lesser extent. In particular, following a 20-min period in which subjects (N = 4) were submitted to a procedure designed to increase the g...

Binocular fixation in the rhesus monkey: spatial and temporal characteristics.

The horizontal and vertical components of the positions of both eyes of rhesus monkeys were measured during periods of binocularly stable eye positions (eye pauses) while the animals fixated a small target. Differences between monocular and binocular viewing, as well as effects of target size and background illumination, were assessed and found to be comparable to similar measures for humans. The scatter of eye position for either eye during binocular viewing had a standard deviation of 6-8 min arc in the horizontal and 7-13 min arc in the vertical meridia. Measurements of vergence and vertical misalignment, taken from binocular positional disparity, showed that for nearly 60% of eye pause time the eyes were misaligned on the fixation target by more than 7 min arc along both horizontal and vertical axes. In addition, the line of gaze during the trial was found to follow certain idiosyncratic tendencies for each monkey, although the positional variability remained relatively constant throughout the fixation trial. These observations suggest that during binocular fusion and stereopsis a mechanism exists that dynamically compensates for the relatively large shifts in retinal image position during fixation.

Binocular retinal image motion during active head rotation

Horizontal binocular eye and head movements of 4 human subjects were recorded by means of the sensor coil-rotating magnetic field technique while they actively rotated their heads about a vertical axis and maintained fixation on a distant target. The frequency and peak-to-peak amplitude of these rotations ranged from about 0.25 Hz to 5 Hz and 30° to 15′. Eye movement compensation of such head rotations was far from perfect and compensation was different in each eye. Average retinal image speed was on the order of 4 deg/sec within each eye and the speed of the changes in retinal image position between the eyes was on the order of 3 deg/sec. Vision, subjectively, remained fused, stable and clear. Attention is called to implications of these results for visual and oculomotor physiology.

An investigation of the mechanisms of eye movement control.

Feedback mechanisms exist in all the periferal sense organs including the eye, which acts as a highly efficient position control servo system. Histological studies so far have not revealed the precise circuitry of the eye movement control system but some information about it can be obtained by a study of the sources of feedback. Existing theories have considered three types of feedback originating in the oculomotor tract, in the proprioceptive fibres of the extrinsic eye muscles and from retinal image displacement. In the present experiments an optical arrangement has been used to vary or eliminate the amount of information available from retinal image motion, and the response of the eye to simple harmonic displacement of a target has been recorded. The response curves of gain (eyeball movement divided by target motion) against frequency indicate that the system is lion linear when the image falls in the retinal region which is insensitive to position. Outside this area, retinal image position is used as negative feedback but the information from the oculomotor tract must be regenerative. There is also evidence for feedback proportional to the first derivative of eyeball position and this function is ascribed to the proprioceptive signals; this form of feedback appears to saturate for large amplitude movements, thus avoiding heavy damping of the flick movements.