Changes In Vergence Research Articles

Dynamics of the accommodation response to abrupt changes in target vergence as a function of age

The dynamic accommodation responses to small, abrupt changes in an accommodation stimulus were studied in two experiments. In the first, responses of 19 subjects with ages distributed between 18 and 49 years were measured for step stimuli of ±1.05 D. In the second, responses to small step stimuli (±0.75, ±1.75 D) were recorded for a group of six ‘young’ (mean age 22, range 16–26 years) and six ‘old’ (mean age 42, range 36–48 years) subjects. In both experiments, the low target vergences always lay within the accommodation range of all subjects. Whether the data were analysed in terms of simple reaction and response times, or in terms of the frequency response, no marked systematic changes with age were found in the dynamics of the response in either of the experiments. It is concluded that, for small stimuli within the amplitude of accommodation, the response dynamics over the adult age range studied remain remarkably constant, even though the amplitude progressively reduces with age as complete presbyopia is approached.

Dynamics of the human linear vestibulo-ocular reflex at medium frequency and modification by short-term training

We study here the effect of a short-term training paradigm on the gain and phase of the human translational VOR (the linear VOR: LVOR). Subjects were exposed to lateral sinusoidal translations on a sled, at 0.5 Hz, 0.3 g peak acceleration. With subjects tracking a remembered target at 1.2 m, the LVOR (slow-phase) under these conditions typically has a phase lead or lag, and a gain that falls short of compensatory. To induce short-term adaptation (training), we presented an earth-fixed visual scene at 1.2 m during sinusoidal translation (x 1 viewing) for 20 minutes, so as to drive the LVOR toward compensatory phase and gain. We examined both the slow-phase and the saccadic responses to these stimuli. Testing after training showed changes in slow-component gain and phase which were mostly but not always in the compensatory direction. These changes were more consistent in naive subjects than in subjects who had previous LVOR experience. Changes in gain were seen with step as well as sinusoidal test stimuli; gain changes were not correlated with vergence changes. There was a strong correlation between gain changes and phase changes across subjects. Fast phases (catch-up saccades) formed a large component of the LVOR under our testing conditions (approximately 30% of the changes in gain but not in phase due to training.

On the linearity of accommodation dynamics

Available experimental data on the gain and phase lags in the closed-loop accommodation responses to stimuli whose dioptric vergence changes sinusoidally with time are examined. It is shown that both phase lags and gain change almost linearly with the temporal frequency of the stimulus. This implies that the phase lags correspond to a frequency-independent time delay and that, if the system is linear, the impulse response should take a symmetrical (sin x/ x) 2 form, with a corresponding standard form of step response. Comparison of this derived step response with the quasi-exponential results found experimentally underlines the limitations of attempting to generalise dynamic accommodation performance from responses obtained with predictable sinusoidally-changing stimuli and the application of a linear model. Consideration of the frequency response which would lead to an exponential step response supports the argument that, although the gain results obtained with sinusoidal stimuli are reasonably representative of those applying with other forms of stimulus, the predictable nature of sinusoidally-changing stimuli makes the derived phase lags inapplicable to non-predictable step or other stimuli. Other limitations in linear models of the response are discussed.

Dynamic accommodation and myopia: §Culhane HM, Winn B. Invest Ophthalmol Vis Sci 1999;40:1968-74

Accommodative effort during nearwork is thought to be a causative factor in the development of myopia. It has been proposed that an anomaly in autonomic control may be a precursor to the development of myopia. In the present study the closed-loop accommodation response after variations in fixation period was investigated in emmetropes, early-onset myopes and late-onset myopes to determine characteristics of reflex accommodation for each refractive group.Closed-loop accommodation responses were measured in a group of emmetropes (n = 7), early-onset myopes (n = 7), and late-onset myopes (n = 7) by use of a dynamic tracking infrared optometer. A variation in fixation period (10 seconds, 60 seconds, and 180 seconds) before an accommodative step was used to stimulate the accommodation control mechanism differentially.Group results of accommodative response times showed that late-onset myopes were significantly affected by the duration of fixation before the change in stimulus vergence. Accommodative response times after 3 minutes of sustained near vision were significantly longer than those observed for other groups for the near-to-far condition. Reaction time appears to be independent of refractive grouping, prior fixation period, and direction of step change.Late-onset myopes showed significantly extended accommodation response times after a sustained near vision task that was demonstrable under well-controlled experimental conditions. The extended response times observed in the present study were consistent with previous reports of refractive shifts in late-onset myopes and early-onset myopes and provide a corollary between reflex and adaptive components of the accommodation response. Potential mechanisms are discussed in an attempt to explain the resultant hysteresis under closed-loop viewing conditions.

The variation of torsion with vergence and elevation

Two recently developed kinematic models of human eye movements predict systematic departures from Listing’s law which are associated with changes in vergence. This vergence-dependent torsion t is proportional to elevation e and vergence v, that is t= kev/2. The proposed value for k is either 1 (Van Rijn, L. J., & Van den Berg, A. V. (1993). Vision Research, 33, 691–708) or 1/2 (Minken, A. W. H., Gielen, C. C. A. M., & Van Gisbergen, J. A. M. (1995). Vision Research, 35, 93–102). One implication of both models is that an eye with a constant fixation direction should exhibit systematic torsional variation during movements of the other eye. This paper therefore examines the torsion produced by moving a fixation target inwards and outwards along the line-of-sight of the right eye at five different viewing elevations (0, ±15 and ±30°). In a monocular analysis, each eye generally showed intorsion during convergence at positive elevation angles, whereas extorsion occurred at negative elevations; the opposite was true during divergence. However, the torsion response was visibly different between the five subjects, and depended on the direction of target motion. In a binocular analysis, cycloversion (mean of left and right eye torsion) varied dramatically both between subjects and between convergence and divergence; however, cyclovergence (torsional difference) was much less variable. Least-squares methods were used to estimate the constant k from monocular torsion, yielding values between 0.2 and 1.0; however, corresponding estimates based on cyclovergence were all close to 1/2. These findings support suggestions that a binocular control system couples the three-dimensional movements of the eyes, and that an existing model of monocular torsion should be generalised to the binocular case.

Perturbation of combined saccade-vergence movements by microstimulation in monkey superior colliculus.

Perturbation of combined saccade-vergence movements by microstimulation in monkey superior colliculus. This study investigated the role of the monkey superior colliculus (SC) in the control of visually (V)-guided combined saccade-vergence movements by assessing the perturbing effects of microstimulation. We elicited an electrical saccade (E) by stimulation (in 20% of trials) in the SC while the monkey was preparing a V-guided movement to a near target. The target was aligned such that E- and V-induced saccades had similar amplitudes but different directions and such that V-induced saccades had a significant vergence component (saccades to a near target). The onset of the E-stimulus was varied from immediately after V-target onset to after V-saccade onset. E-control trials, where stimulation was applied during fixation of a V-target, yielded the expected saccade but no vergence. By contrast, early perturbation trials, where the E-stimulus was applied soon after the onset of the V-target, caused an E-triggered response with a clear vergence component toward the V-target. Midflight perturbation, timed to occur just after the monkey initiated the movement toward the target, markedly curtailed the ongoing vergence component during the saccade. Examination of pooled responses from both types of perturbation trials showed weighted-averaging effects between E- and V-stimuli in both saccade and fast vergence components. Both components exhibited a progression from E- to V-dominance as the E-stimulus was delayed further. This study shows that artificial intervention in the SC, while a three-dimensional (3D) refixation is being prepared or is ongoing, can affect the timing (WHEN) and the metric specification (WHERE) of both saccades and vergence. To explain this we interpret the absence of overt vergence in the E-controls as being caused by a zero-vergence change command rather than reflecting the mere absence of a collicular vergence signal. In the perturbation trials, the E-evoked zero-vergence signal competes with the V-initiated saccade-vergence signal, thereby giving rise to a compromised 3D response. This effect would be expected if the population of movement cells at each SC site is tuned in 3D, combining the well-known topographical code for direction and amplitude with a nontopographical depth representation. On E-stimulation, the local population would yield a net saccade signal caused by the topography, but the cells coding for different depths would be excited equally, causing the vergence change to be zero.

Response AC/A ratios are elevated in myopic children

In children little is known about the relationship between the AC/A ratio and the development of myopia, although they have been linked in adults. The purpose of this study was to investigate the interaction between accommodation and convergence and its relationship to refractive errors in children. Accommodation was measured for the right eye using the Canon R-1 autorefractor, and concomitant changes in vergence were assessed using a Maddox rod and a Risley prism before the left eye. Thirty-three myopic and 68 emmetropic children were tested wearing best subjective correction while looking at a distant (4.0m) letter array and a near (0.33m) one through additional plus and minus lenses. Lens-induced and distance-induced response AC/A ratios were calculated from the data. Both types of AC/A ratios are elevated in myopic children, who show reduced accommodation and enhanced accommodative convergence. Myopic children with esophoria underaccommodate at near. This suggests that a child who is esophoric must relax accommodation to reduce accommodative convergence and maintain single binocular vision. The reduction in accommodation could produce blur during near work, which could induce myopia as in animal models.

Response AC/A ratios are elevated in myopic children.

In children little is known about the relationship between the AC/A ratio and the development of myopia, although they have been linked in adults. The purpose of this study was to investigate the interaction between accommodation and convergence and its relationship to refractive errors in children. Accommodation was measured for the right eye using the Canon R-1 autorefractor, and concomitant changes in vergence were assessed using a Maddox rod and a Risley prism before the left eye. Thirty-three myopic and 68 emmetropic children were tested wearing best subjective correction while looking at a distant (4.0 m) letter array and a near (0.33 m) one through additional plus and minus lenses. Lens-induced and distance-induced response AC/A ratios were calculated from the data. Both types of AC/A ratios are elevated in myopic children, who show reduced accommodation and enhanced accommodative convergence. Myopic children with esophoria underaccommodate at near. This suggests that a child who is esophoric must relax accommodation to reduce accommodative convergence and maintain single binocular vision. The reduction in accommodation could produce blur during near work, which could induce myopia as in animal models.

Short-latency vergence eye movements induced by radial optic flow in humans: dependence on ambient vergence level.

Radial patterns of optic flow, such as those experienced by moving observers who look in the direction of heading, evoke vergence eye movements at short latency. We have investigated the dependence of these responses on the ambient vergence level. Human subjects faced a large tangent screen onto which two identical random-dot patterns were back-projected. A system of crossed polarizers ensured that each eye saw only one of the patterns, with mirror galvanometers to control the horizontal positions of the images and hence the vergence angle between the two eyes. After converging the subject's eyes at one of several distances ranging from 16.7 cm to infinity, both patterns were replaced with new ones (using a system of shutters and two additional projectors) so as to simulate the radial flow associated with a sudden 4% change in viewing distance with the focus of expansion/contraction imaged in or very near both foveas. Radial-flow steps induced transient vergence at latencies of 80-100 ms, expansions causing increases in convergence and contractions the converse. Based on the change in vergence 90-140 ms after the onset of the steps, responses were proportional to the preexisting vergence angle (and hence would be expected to be inversely proportional to viewing distance under normal conditions). We suggest that this property assists the observer who wants to fixate ahead while passing through a visually cluttered area (e.g., a forest) and so wants to avoid making vergence responses to the optic flow created by the nearby objects in the periphery.

The effects of eccentricity and vergence angle upon the relative tilt of corresponding vertical and horizontal meridia revealed using the minimum motion paradigm.

When the corresponding horizontal meridia of the two eyes are aligned, the corresponding vertical meridia are tilted outwards in a temporal direction, a phenomenon first described by Helmholtz. However, it is not known if this effect is confined to the principal meridia or whether the same relationship exists between corresponding horizontal and corresponding vertical meridia at eccentric retinal locations. We sought to address this issue by exploiting the technique of Nakayama (1977 Proceedings of the Society of Photo-Optical Instrument Engineers 120 2-9) in which the positions of alternating dichoptic images that produce minimal apparent motion were used to measure the relative tilt of corresponding meridia at a range of eccentricities up to +/- 16 deg away from the fovea. Stimuli were composed of dichoptic images, one containing a blank field and the other a pair of dots, which alternated at a rate of 0.63 Hz and the relative tilt (binocular orientation difference) between the pairs of dots presented to the two eyes was varied between +/- 11 degrees. Nonius lines were used to maintain vergence angle, which was varied between 28 cm and infinity. Subjects judged which pair of alternating images produced the smallest amount of apparent motion (position change). It was found that at all eccentricities examined the corresponding horizontal meridia were generally aligned but the corresponding vertical meridia were consistently offset (extorted) by about +/- 2 degrees. The tilts of corresponding principal meridia were typically unaffected when vergence angle was varied, indicating that little or no cyclovergence accompanied changes in horizontal vergence. The results suggest that the binocular correspondence system appears to be mapped by a horizontal shear distortion that extends to retinal locations at least as far as 16 deg away from the foveae. The invariant extortion of corresponding principal vertical meridia with vergence state is consistent with previous suggestions that the empirical vertical horopter becomes progressively inclined with respect to the vertical as viewing distance increases.

Monocularly programmed human saccades during vergence changes?

1. When binocular fixation is shifted to a new target located at a different distance and in a different direction from initial fixation, a binocularly unbalanced saccade occurs at or near the onset of the composite eye movement. Those saccades typically produce good post-saccadic foveation of the target by one eye or the other. 2. Following such saccades, the better-aligned eye is typically as well aimed at the target as after pure versional saccades, but the partner eye deviates much more, thus requiring asymmetrical post-saccadic vergence movement. 3. This phenomenon suggests that during binocular viewing, the retinal eccentricity of a new-target's image from one of the eyes can be used to programme that eye's own saccade, so that it arrives reliably on target; and that the images of that target from both eyes participate in generating the saccadic excursion of the partner eye. 4. The ecologically useful result is rapid achievement of a high-resolution monocular view of the new target, although full binocular foveation is achieved much later.

Ocular vergence measurement in projected and collimated simulator displays.

The purpose of this study was to investigate electrooculography (EOG) as a measurement of ocular vergence in both collimated and projected simulator environments. The task required participants to shift their gaze between a central fixation point and a target appearing at one of three eccentricities. EOG was effective in recording ocular vergence. The EOG results were similar between collimated and projected displays, except for differences in vergence changes during lateral movement of the eyes, and ocular excursions downward elicited a greater EOG response than the reverse upward movement. The computer-based technique of recording vergence was found to produce measurable traces from a majority of participants. The technique has potential for further development as a tool for measuring ocular vergence in virtual environments where methods that require the wearing of head-mounted apparatus to track ocular structures (e.g., the pupil), which cannot be worn at the same time as a flight or flight-simulator helmet, are unsuitable.

Relationships among visual acuity demands, convergence, and nystagmus in patients with manifest/latent nystagmus

Background: We investigated the role convergence plays in nystagmus dampening, in particular, relationships among visual acuity demands, convergence, and nystagmus. Previously we showed that subjects with idiopathic infantile nystagmus exhibit a range of responses to acuity targets, one of which is nystagmus blockage syndrome. We report herein eye movement responses to acuity targets of patients with manifest/latent nystagmus. Methods: Fourteen patients, 11 with latent or manifest latent nystagmus and 3 with combined manifest latent with infantile nystagmus, were asked to indicate the direction of the gap in Landolt C optotypes while their eye movements were recorded. Results: The tested patients exhibited various responses to acuity demands: (1) dampening of nystagmus with convergence (i.e., nystagmus blockage syndrome) ( 5 14 patients), (2) changes in vergence without nystagmus dampening (2 patients), (3) decrease of nystagmus without convergence (2 patients), and (4) little change in nystagmus or vergence (5 patients). In nystagmus blockage syndrome the amount of convergence increased with acuity demands in two of five patients and the convergence duration in four of five patients; nystagmus dampening increased with acuity demands in one of five patients and the blockage duration in four of five patients. Conclusions: Many, but not all, patients with manifest/latent nystagmus, similar to those with infantile nystagmus, used convergence to dampen their nystagmus. The convergence response tended to increase with acuity demands, but the amount of dampening was idiosyncratic and not predictably related to the measured convergence across patients.

Visual-vestibular interaction with vergence

Introduction. Vergence, or fixation distance, influences the vestibulo-ocular reflex (VOR). The VOR-gain increases with decreasing fixation distance. This distance is primarily resolved visually. Is visual control maintained at high frequency head movements? We investigated whether the visual system, during high frequency headshaking and a quick change of gaze direction, controls the eye movements in such a way that binocular vision is maintained. Material and methods. Two-dimensional eye movements were recorded with the scleral coil technique during active head rotation (0.5–4 Hz) while fixating an object. The subject was wearing glasses containing two displacement prisms (OD = 4D, OS = 1D). During headshaking, the head was tilted whilst the subject looked through the prisms at the fixation object. Results. Within 1.5 s after head tilt, the correct amount of vergence change was reached. For high shake frequencies this occurred within 0.5 s. The vergence amplitudes before and after tilt were comparable. Conclusion. It appears that the eyes can compensate for head movements independent of gaze direction even at high shake frequencies. The VOR controls the high frequency eye movements and the visual system updates at a lower frequency mainly when the head velocity is almost zero.

Otolith and Semicircular Canal Contributions to the Human Binocular Response to Roll Oscillation

Three normal human subjects were oscillated about their naso-occipital axis in a supine position at 0.4 Hz and 0.1 Hz, both in darkness and in the light with a structured fixation target. The same subjects were oscillated in roll about an upright position, at the same frequencies, in darkness; and also about axes directed 20° and 40° to the left and to the right of the midsagittal plane, at 0.4 Hz, in darkness. Three-dimensional binocular eye movements were recorded using video-oculography. All stimuli induced a predominantly torsional nystagmus with small disconjugate head-vertical (skew) and conjugate head-horizontal components. For roll oscillation, the torsional slow phase velocity gain was higher in the light and generally increased with the stimulation frequency. In darkness, only one subject had significantly higher torsional gains in the upright compared to the supine position (12% difference), suggesting that the otolith contribution to the roll response is minimal at the frequencies tested. The slow phase velocity gain of the skew increased with fixation in two subjects, but decreased in the third subject; these changes were related to changes in eye vergence. In the upright position, with oblique axes of rotation, the head-vertical eye movements were asymmetrical, with the outermost eye executing a larger amplitude movement. The disconjugate head-vertical eye movements observed can be explained by the pattern of vertical semicircular canal stimulation and their connections to the extraocular muscles. In humans, skewing of the eyes may compensate for the eccentricity of the foveae which lie in the temporal retina and undergo vertical translations in opposite directions when the eyes tort.

Looking at the task in hand: vergence eye movements and perceived size.

A retinal afterimage of the hand changes size when the same unseen hand is moved backwards and forwards in darkness. We demonstrate that arm movements per se are not sufficient to cause a size change and that vergence eye movements are a necessary and sufficient condition for the presence of the illusory size change. We review previous literature to illustrate that changing limb position in the dark alters vergence angle and we explain the illusion via this mechanism. A discussion is provided on why altering limb position causes a change in vergence and we speculate on the underlying mechanisms.

A cross-section through the Zilair Nappe (southern Urals)

A structural transect in the Zilair-Kugarchi area involves the western part of the Suvanyak Complex, the Zilair Nappe and the eastern part of the foreland thrust and fold belt. This section has been analyzed using field, microstructural and seismic data. The cross-section shows the transition from the hinterland to the foreland in the footwall to the suture of the southern Urals. The rocks involved range from early Palaeozoic to Permian in age. A characteristic of the Zilair Nappe is the dominance of a succession of volcanic greywackes and mudrocks of Late Devonian age (Zilair Formation). The metamorphic grade decreases from east to west, from greenschist facies to diagenetic conditions. The structure of the cross-section mainly comprises west-directed thrusts and thrust-related folds with an associated cleavage. Fold vergence changes along the section depending on of the distance to the associated thrust and its geometry. The Zilair thrust which separates the Zilair Nappe from the foreland thrust and fold belt accommodated ca. 10 km displacement and the characteristics of the deformation are similar on both sides of it. The contact between the Zilair Nappe and Suvanyak Complex is a west-dipping normal fault that does not represent a major tectonic boundary.

Spatiotopic cues enhance the dynamic accommodation response profile

Purpose: To determine the influence of optically-induced and real changes in stimulus vergence on the profile of the accommodation response. Method: Accommodation was measured on 8 visually normal subjects (mean age 25.6±6.0 years) using a dynamic tracking infra-red optometer. A Badal stimulus optometer was used to induce optical changes in stimulus vergence which eliminated spatiotopic cues. Targets were also located in real space to provide both retinotopic and spatiotopic cues. Step changes in stimulus vergence were modulated over a 2 D (2–4 D) range for both conditions. Data were collected at a sampling rate of 102.4 Hz using a digital storage oscilloscope which was interfaced to an IBM PC. Results: Mean responses are presented below. An ANOVA revealed no significant difference between overall response times to optically-induced and real changes in stimulus vergence although the response profiles were different between conditions. Conclusions: Spatiotopic cues enhance the accommodation response profile for step changes in stimulus vergence by reducing initial directional error and ensuring the magnitude of the response is appropriate to the stimulus demand. However, overall response times were found to be similar under both viewing conditions.

Spatiotopic cues enhance the dynamic accommodation response profile

Purpose: To determine the influence of optically‐induced and real changes in stimulus vergence on the profile of the accommodation response.Method: Accommodation was measured on 8 visually normal subjects (mean age 25.6±6.0 years) using a dynamic tracking infra‐red optometer. A Badal stimulus optometer was used to induce optical changes in stimulus vergence which eliminated spatiotopic cues. Targets were also located in real space to provide both retinotopic and spatiotopic cues. Step changes in stimulus vergence were modulated over a 2 D (2–4 D) range for both conditions. Data were collected at a sampling rate of 102.4 Hz using a digital storage oscilloscope which was interfaced to an IBM PC.Results: Mean responses are presented below.An ANOVA revealed no significant difference between overall response times to optically‐induced and real changes in stimulus vergence although the response profiles were different between conditions.Conclusions: Spatiotopic cues enhance the accommodation response profile for step changes in stimulus vergence by reducing initial directional error and ensuring the magnitude of the response is appropriate to the stimulus demand. However, overall response times were found to be similar under both viewing conditions.

Modulation of vergence by off-vertical yaw axis rotation in the monkey: normal characteristics and effects of space flight.

Horizontal movements of both eyes were recorded simultaneously using scleral search coils in 2 rhesus monkeys before and after the COSMOS 2229 space-flight of 1992-1993. Another 9 monkeys were tested at comparable time intervals and served as controls. Ocular vergence, defined as the difference in horizontal position between the left and right eyes, was measured during off-vertical yaw axis rotation (OVAR) in darkness. Vergence was modulated sinusoidally as a function of head position with regard to gravity during OVAR. The amplitude of peak-to-peak modulation increased with increments in tilt of the angle of the rotational axis (OVAR tilt angle) that ranged from 15 degrees to 90 degrees. Of the 11 monkeys tested, 1 had no measurable modulation in vergence. In the other 10, the mean amplitude of the peak to peak modulation was 5.5 degrees +/- 1.3 degrees at 90 degrees tilt. Each of these monkeys had maximal vergence when its nose was pointed close to upward (gravity back; mean phase: -0.9 degree +/- 26 degrees). After space flight, the modulation in vergence was reduced by over 50% for the two flight monkeys, but the phase of vergence modulation was not altered. The reduction in vergence modulation was sustained for the 11-day postflight testing period. We conclude that changes in vergence are induced in monkeys by the sinusoidal component of gravity acting along the naso-occipital axis during yaw axis OVAR, and that the modulation of the vergence reflex is significantly less sensitive to linear acceleration after space flight.