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Biomedical subjects

F A Miles

Publications and source records attributed to F A Miles.

At least 19 recordsLinked to original sources
PubMed1992

Ocular compensation for self-motion. Visual mechanisms.

In monkeys, there are several reflexes that generate eye movements to compensate for the observer's own movements. Two vestibuloocular reflexes compensate selectively for rotational (RVOR) and translational (TVOR) disturbances of the head, receiving their inputs from the semicircular canals and otolith organs, respectively. Two independent visual tracking systems that deal with residual disturbances of gaze are manifest in the two components of the optokinetic response: the indirect or delayed component (OKNd) and the direct or early component (OKNe). We hypothesize that OKNd--like the RVOR--is phylogenetically old, being found in all animals with mobile eyes, and that it evolved as a backup to the RVOR to compensate for rotational disturbances of gaze. Indeed, optically induced changes in the gain of the RVOR result in parallel changes in the gain of OKNd, consistent with the idea of shared pathways as well as shared functions. In contrast, OKNe--like the TVOR--seems to have evolved much more recently in frontal-eyed animals and, we suggest, acts as a backup to the TVOR to deal primarily with translational disturbances of gaze. Frontal-eyed animals with good binocular vision must be able to keep both eyes directed at the object of regard irrespective of proximity, and in order to achieve this during translational disturbances, the output of the TVOR is modulated inversely with the viewing distance. OKNe shares this sensitivity to absolute depth, consistent with the idea that it is synergistic with the TVOR and shares some of its central pathways. There is evidence that OKNe is also sensitive to relative depth cues such as motion parallax, which we suggest helps the system to segregate the object of regard from other elements in the scene. However, there are occasions when the global optic flow cannot be resolved into a single vector useful to the oculomotor system (e.g., when the moving observer looks towards the direction of heading). We suggest that on such occasions a third independent tracking mechanism, the smooth pursuit system, is deployed to stabilize gaze on the local feature of interest. In this scheme, the pursuit system has an attentional focusing mechanism that spatially filters the visual motion inputs driving the oculomotor system. The major distinguishing features of the 3 visual tracking mechanisms are summarized in Table 1.

Animals↗
PubMed1992

Effects of stationary textured backgrounds on the initiation of pursuit eye movements in monkeys.

1. The initial ocular pursuit of small target spots (0.25 degrees diam) that suddenly start to move at constant speed (ramps) was recorded in four rhesus monkeys with the electromagnetic search coil technique. All target motions were horizontal, and both eyes were monitored. 2. In agreement with the observations of Keller and Khan, stationary textured backgrounds substantially reduced the initial eye acceleration achieved during pursuit but did not affect its latency. Correlation techniques were used to assess the changes in the eye speed profiles and indicated that the reduction in eye acceleration due to the background was a linear function of the logarithm of target speed over the range investigated (5-40 degrees/s), averaging 60% with the fastest targets. 3. Selectively excluding the background texture from the path of the target with a horizontal strip of card (vertical width, 4 degrees) reduced the impact of the background only slightly, and, even when the vertical width of the card was increased to 60 degrees, the effect of the background was not entirely eliminated. Thus the effect involves regions of the visual field well beyond the target and is not due simply to the reduced physical salience (contrast) of the target spot. Such spatially remote interactions suggest that the neurons decoding the target's motion have very extensive visual receptive fields. 4. Textured backgrounds also caused similar reductions in the eye acceleration during initial pursuit when, before the ramps, the fixated target spots stepped forward, i.e., stepped in the direction of the subsequent ramps (step ramps). In this situation, as with no steps, initial target ramps were foveofugal. When the fixated target spots were stepped back before moving forward so that initial target ramps were foveopetal, textured backgrounds now also delayed the onset of pursuit, and the reductions in eye acceleration were not seen until some time later when tracking resulted from foveofugal target-ramp motion. Selectively excluding the texture from the path of the target with a narrow strip of card eliminated any delays in the onset of pursuit to step ramps, but the later reductions in eye acceleration were still evident. These step-ramp data indicate that the mechanisms decoding foveofugal and foveopetal target ramps differ markedly in their sensitivity to textured backgrounds. That backgrounds can influence the latency and the initial eye acceleration independently is consistent with the idea that there are independent trigger and drive mechanisms for the decoding of target motions.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1991

Ocular responses to translation and their dependence on viewing distance. I. Motion of the observer.

1. The horizontal eye movements induced by acceleration along the interaural axis were recorded from five monkeys (Macaca mulatta) by the use of the electromagnetic search-coil technique. Animals sat on a sled that was moved briefly in darkness along a linear track (bell-shaped acceleration profile: peak, 400 mm/s2; duration, 200 ms). Immediately before acceleration, animals fixated one of five target lights located at distances ranging from 16 to 150 cm. During fixation, the horizontal positions of both eyes were used to check vergence, while accommodation was monitored with an infrared optometer. 2. Sled motion induced eye movements that were generally smooth and compensatory, e.g., rightward sled motion elicited leftward eye movement. We attribute these responses to a translational vestibuloocular reflex (TVOR) that senses the sled motion through the otolith organs. However, in three animals, these responses were preceded by weak anticompensatory movements (duration, less than 40 ms; amplitude, less than 10% of the maximum compensatory response). 3. Geometry indicates that, during brief sled motion, the eye movements required to keep gaze aligned on a particular location ("full" compensation) are inversely proportional to the viewing distance. Response measures based on the computed eye velocity profiles, such as the velocity achieved at specific times ("time slices") or the peak values of the estimated covariance functions, all indicated that compensatory responses were a linear function of the inverse of the prior viewing distance. Cross-correlation analyses indicated that the effect of the prior viewing distance was to scale responses, although detailed spectral analyses revealed that high-frequency components (greater than 10 Hz) tended to scale less vigorously than lower ones. 4. The adequacy of the compensatory eye movements was assessed by calculating the gain (response recorded/response required for full compensation). Regardless of the response measure used, gains varied considerably from one animal to another and, in some particular animals, from one direction to another but showed a general tendency to increase with viewing distance. For example, on the basis of the peak eye velocity achieved within 250 ms of the onset of sled motion, mean gain at 16 cm was 0.74 (range, 0.48-1.01), whereas at 150 cm it was 1.25 (range, 0.67-1.73). 5. Using wedge prisms to dissociate vergence and accommodation indicated that ocular responses to sled motion were sensitive to selective increases in either vergence (base-out prism with the most distant target) or accommodation (base-in prism with the nearest target). However, the magnitude of the effects showed considerable variability from one animal to another and, in some particular animals, from one direction to another.(ABSTRACT TRUNCATED AT 400 WORDS)

Acceleration↗
PubMed1991

Ocular responses to translation and their dependence on viewing distance. II. Motion of the scene.

1. The ocular following responses induced by brief (100-ms) movements of the visual scene were examined for their dependence on viewing distance in 5 monkeys (Macaca mulatta). The horizontal positions of both eyes and the vertical position of one eye were recorded using the electromagnetic search-coil technique. Accommodation was monitored in selected experiments by use of an infrared optometer. Test patterns (random dots) were back-projected onto a translucent tangent screen facing the animal. Six viewing distances were used (range, 20-150 cm), the size and speed of the image on the screen being adjusted for each so as to preserve a constant retinal image. 2. Response measures based on the amplitude of the first peak in the eye acceleration profile or the eye velocity achieved at specific times all indicated that ocular following responses were inversely related to viewing distance, the relationship being linear for the earliest measures. On average, the sensitivity to viewing distance was comparable with that reported for the translational vestibuloocular reflex (TVOR) in the preceding paper: as viewing distance increased from 20 cm, ocular following decremented at a mean rate (+/- SD) of 17 +/- 3% per m-1, while the TVOR decremented at a mean rate (+/- SD) of 18 +/- 1% per m-1. 3. Ocular following responses showed the postsaccadic enhancement described by Kawano and Miles regardless of viewing distance. To a first approximation, the effects of postsaccadic enhancement and viewing distance summed linearly. 4. The dependence of ocular following on speed showed the progressive saturation previously described by Miles et al. at all viewing distances, the peak eye velocity achieved being inversely related to the viewing distance, indicating that the saturation must originate upstream of the dependence on viewing distance. Under normal viewing conditions, this speed saturation will tend to offset the dependence on viewing distance because the retinal slip speeds experienced by the moving observer will tend to vary inversely with viewing distance, resulting in greater saturation with nearer viewing. 5. Wedge prisms were used to dissociate vergence and accommodation and indicated that ocular following responses were sensitive to selective increases in either vergence (base-out prism with the screen at 100 cm) or accommodation (base-in prism with the screen at 20 cm). However, as with the TVOR, the magnitude of the effects showed considerable variability from one animal to another and, in some particular animals, from one direction to another.(ABSTRACT TRUNCATED AT 400 WORDS)

Acceleration↗
PubMed1990

Local ocular compensation for imposed local refractive error.

Chicks were raised in a low-ceiling environment to find out if their eye growth could compensate for locally imposed hyperopic refractive errors. These chicks became selectively more myopic in the upper visual field than chicks raised in a high-ceiling environment. The vitreous chamber in the low-ceiling birds showed a selective elongation in the ventral region that was not seen in the eyes of the high-ceiling birds. This morphological difference was small, but probably adequate to account for the additional myopia in the low-ceiling birds. These results are consistent with the idea of a visually mediated growth mechanism regulating the local refractive state across the entire visual field so that it matches the customary viewing conditions. Such a mechanism might account for the finding of Fitzke, Hayes, Hodos, Holden and Low (Journal of Physiology, London, 369, 33-44, 1985) that the refractive errors in the lower field are exactly appropriate for focusing the image of the ground on the retina.

Animals↗
PubMed1990

Short latency ocular-following responses in man.

The ocular-following responses elicited by brief unexpected movements of the visual scene were studied in human subjects. Response latencies varied with the type of stimulus and decreased systematically with increasing stimulus speed but, unlike those of monkeys, were not solely determined by the temporal frequency generated by sine-wave stimuli. Minimum latencies (70-75 ms) were considerably shorter than those reported for other visually driven eye movements. The magnitude of the responses to sine-wave stimuli changed markedly with stimulus speed and only slightly with spatial frequency over the ranges used. When normalized with respect to spatial frequency, all responses shared the same dependence on temporal frequency (band-pass characteristics with a peak at 16 Hz), indicating that temporal frequency, rather than speed per se, was the limiting factor over the entire range examined. This suggests that the underlying motion detectors respond to the local changes in luminance associated with the motion of the scene. Movements of the scene in the immediate wake of a saccadic eye movement were on average twice as effective as movements 600 ms later: post-saccadic enhancement. Less enhancement was seen in the wake of saccade-like shifts of the scene, which themselves elicited weak ocular following, something not seen in the wake of real saccades. We suggest that there are central mechanisms that, on the one hand, prevent the ocular-following system from tracking the visual disturbances created by saccades but, on the other, promote tracking of any subsequent disturbance and thereby help to suppress post-saccadic drift. Partitioning the visual scene into central and peripheral regions revealed that motion in the periphery can exert a weak modulatory influence on ocular-following responses resulting from motion at the center. We suggest that this may help the moving observer to stabilize his/her eyes on nearby stationary objects.

Humans↗
PubMed1989

Ocular responses to linear motion are inversely proportional to viewing distance.

Eye movements exist to improve vision, in part by preventing excessive retinal image slip. A major threat to the stability of the retinal image comes from the observer's own movement, and there are visual and vestibular reflexes that operate to meet this challenge by generating compensatory eye movements. The ocular responses to translational disturbances of the observer and of the scene were recorded from monkeys. The associated vestibular and visual responses were both linearly dependent on the inverse of the viewing distance. Such dependence on proximity is appropriate for the vestibular reflex, which must transform signals from Cartesian to polar coordinates, but not for the visual reflex, which operates entirely in polar coordinates. However, such shared proximity effects in the visual reflex could compensate for known intrinsic limitations that would otherwise compromise performance at near viewing.

Animals↗
PubMed1987

Optically induced changes in the couplings between vergence and accommodation.

Vergence-induced accommodation (V-A) and accommodation-induced vergence (A-V) were measured in human subjects before and after they had worn various optical devices for 30 min. Laterally displacing periscopic spectacles, which increase the required change in vergence per unit change in accommodation, caused decreases and increases in the gain of V-A and A-V responses, respectively. These observations are consistent with the view that the gain of the neural cross-linkages between vergence and accommodation are subject to adaptive regulation. However, there were strong asymmetries: Medially displacing periscopes (cyclopean spectacles), which reduce the required change in vergence per unit change in accommodation to zero, were almost totally without effect. Base-out prisms, which increase the required convergence by an amount that is constant for all viewing distances, caused downward and upward shifts in the V-A and A-V response curves, respectively. Base-in prisms, which reduce the required convergence by a constant amount for all viewing distances, caused downward shifts of A-V curves but had no significant effect on V-A curves. These effects of prisms are in essential agreement with the work of others and confirm the existence of adaptive elements that regulate the bias in the vergence and/or accommodation control systems. Secondary effects of wearing periscopes and prisms indicated a certain lack of specificity in the sensing of gain and bias errors: vertical shifts of V-A and A-V curves (resembling those seen with base-out prisms) often occurred with the laterally displacing periscopes, and gain changes (generally resembling those seen with laterally displacing periscopes) often occurred with the base-out prisms.

Accommodation, Ocular↗
PubMed1986

Floccular lesions abolish adaptive control of post-saccadic ocular drift in primates.

After several days of exposure to optically-imposed post-saccadic retinal slip, the saccades of normal monkeys acquire an exponential ocular drift. This drift is in the direction of the imposed image motion, and persists in the dark. It has been argued that these changes result from the operation of a visually mediated adaptive mechanism that normally functions to minimize post-saccadic ocular drift. Adaptation to persistent post-saccadic retinal slip was assessed in two rhesus monkeys before and after bilateral ablations of the flocculi and portions of the paraflocculi ("flocculectomy"). After flocculectomy, both monkeys showed some post-saccadic ocular drift. Flocculectomized animals also failed to adapt to optically-imposed post-saccadic slip. We infer from this that the flocculi and/or paraflocculi are necessary for the successful suppression of post-saccadic ocular drift.

Adaptation, Ocular↗
PubMed1986

Short-latency ocular following responses of monkey. I. Dependence on temporospatial properties of visual input.

The ocular following responses elicited by brief unexpected movements of the visual scene were studied in 10 rhesus monkeys. Test patterns were either random dots or sine-wave gratings [spatial frequency (Fs) 0.046-1.06 cycles per degree (c/degree)]. Test stimuli were velocity steps [speed (V) 5-400 degrees/s] of 100-ms duration, applied 50 ms after spontaneous saccades to avoid saccadic intrusions. Eye velocity response profiles were nonmonotonic and idiosyncratic, but consistent and closely time-locked to stimulus onset. Two measures of response amplitude were used: initial peak in eye velocity (ei), and average final eye velocity over the period of 110-140 ms measured from stimulus onset (ef). Using random dot patterns, response latencies were short, e.g., when the criterion for onset was an eye acceleration of 100 degrees/s2, mean latency (+/- SE) for eight monkeys with a 40 degrees/s test ramp was 51.5 +/- 0.6 ms. Using gratings of low spatial frequency (Fs less than 0.5 c/degree), latency was inversely related to, and solely a function of, contrast and temporal frequency, Ft (where Ft = V X Fs). We conclude from the latter that ocular following is triggered by local changes in luminance, and propose a model of the detection mechanism that reproduces all the essential features of these data. Moderate low-pass spatial filtering ("blurring") of the random dot pattern, by interposing a sheet of ground glass between the animal and the scene, progressively increased the response latency and decreased ef, but ei was either little affected or increased. When used with gratings, the ground glass simply reduced the contrast (range: 0.5-0.003), with very similar consequences for ocular following: latency increased and ef decreased, but ei changed little over the first decade of contrast reduction, increased over the second, and began to show attenuation (often pronounced) only at the lowest contrast. We suggest that these anomalous increases in ei with reductions in contrast are secondary to the delay in response onset and might be explained if the motion detectors responsible for triggering ocular following act as a gate for integrated retinal slip inputs to the tracking system proper: the delay in detection causes a buildup in the error signal driving the tracking response. En masse movement of the visual field was not the optimal stimulus for ocular following.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1986

Short-latency ocular following responses of monkey. II. Dependence on a prior saccadic eye movement.

The ocular following responses elicited by brief unexpected movements of the visual scene were studied in eight rhesus monkeys. Test patterns were random dots except in one experiment when sine-wave gratings were used. Test stimuli were velocity steps of 100-ms duration applied after spontaneous saccades. Two response measures were used: the initial peak in the eye velocity profile (ei), and the average final eye velocity over the period of 110-140 ms measured from stimulus onset (ef). Responses were best when the test ramps began soon after saccades and attenuated progressively as the postsaccadic delay interval was increased: postsaccadic enhancement of ocular following. The decline in ei was roughly exponential: average time constant, 60 ms; average asymptote, 22%. Later measures (ef) were generally less affected. We suggest that this transient enhancement aids the visual suppression of postsaccadic ocular drifts (glissades) and the tracking of moving images newly acquired with a saccade. The magnitude of the postsaccadic enhancement was dependent on the amount of retinal stimulation during the antecedent saccade; when this stimulation was compromised, as when a vertical saccade was made while viewing a grating pattern with vertically oriented stripes, subsequent enhancement of ocular following was much reduced. Further, saccade-like conditioning movements of the visual scene resulted in an enhancement of the ocular following, elicited by subsequent test ramps, that was similar in magnitude and time course to that in the wake of real saccades. We conclude that the postsaccadic enhancement of ocular following is largely due to the visual stimulation produced by the saccade sweeping the scene across the retina. Data obtained with the visual field partitioned into central and peripheral regions (center 20-60 degrees diam) and with gaze centered suggested that the short-latency ocular following system and the enhancement mechanism that modulates it both receive their major inputs from the central 40 degrees of the retina. Further, when this central region was partitioned, enhancement was obtained only when the conditioning and test stimuli were presented to the same region of retina. Visual enhancement showed only weak interocular transfer: the conditioning and test stimuli had to be seen by the same eye to produce appreciable enhancement. These data suggest that the enhancement involves local spatial interactions at an "early" point in the visual pathway before the inputs from the two eyes have converged. When the conditioning and test stimuli impinged on different regions of the retina, brief powerful suppression of ocular following was obtained.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1986

Short-latency ocular following responses of monkey. III. Plasticity.

The adaptability of the ocular following responses elicited by brief movements of the visual scene was studied in five rhesus monkeys. Adapting stimuli were double-ramp sequences, designed to initiate ocular following and then induce consistent visual errors that would challenge any adaptive mechanism regulating the performance of such tracking. Both ramps lasted 150 ms and could differ from one another in speed (speed steps) or direction (direction steps), depending on the experiment. Monkeys were repeatedly exposed to these adapting stimuli over a period of 3 days with overnight rest periods in darkness. Ocular following responses were calibrated with 100-ms test ramps that covered a range of speeds (10-100 degrees/s) and directions (right, left, up, down) and were applied 50 ms after spontaneous saccades. Repeated exposure to speed steps involving abrupt increases or decreases in speed resulted in clear increases or decreases, respectively, in the ocular following responses elicited by the standard test ramps. Changes were greatest during the first part of the day, with partial recovery during the overnight periods. Changes were specific to the direction and, to a lesser extent, the speed of the adapting stimuli. Repeated exposure to direction steps involving abrupt 90 degrees counterclockwise changes in the direction of movement resulted in the emergence of a counterclockwise orthogonal component of ocular following. Orthogonal responses differed from isogonal ones in having smoother response profiles and in showing no overnight recovery. All of the recorded changes were adaptive insofar as they would result in improved tracking of the second ramp in the double-ramp sequence. It is concluded that the ocular following responses of the monkey are subject to extensive visually mediated adaptive regulation that influences both the magnitude and direction of tracking.

Adaptation, Ocular↗
PubMed1985

Changes in the coupling between accommodation and vergence eye movements induced in human subjects by altering the effective interocular separation.

It has usually been thought that the coupling between accommodation and convergence of the eyes is fixed and not modifiable by experience. Experiments are reported which show that the ratio of accommodative vergence to the accommodation stimulus, the stimulus AC/A ratio (one measure of the coupling), is elevated by brief periods (approximately 30 min) of experience of viewing the world through periscopic spectacles which increase the effective interocular separation. Experience of viewing through 'cyclopean' spectacles, which superimpose the line of sight of the two eyes, reduced the stimulus AC/A ratio in one subject and increased it in another, while it remained hardly affected in a third.

Accommodation, Ocular↗
PubMed1985

Visually induced adaptive changes in primate saccadic oculomotor control signals.

Saccades are the rapid eye movements used to change visual fixation. Normal saccades end abruptly with very little postsaccadic ocular drift, but acute ocular motor deficits can cause the eyes to drift appreciably after a saccade. Previous studies in both patients and monkeys with peripheral ocular motor deficits have demonstrated that the brain can suppress such postsaccadic drifts. Ocular drift might be suppressed in response to visual and/or proprioceptive feedback of position and/or velocity errors. This study attempts to characterize the adaptive mechanism for suppression of postsaccadic drift. The responses of seven rhesus monkeys were studied to postsaccadic retinal slip induced by horizontal exponential movements of a full-field stimulus. After several hours of saccade-related retinal image slip, the eye movements of the monkeys developed a zero-latency, compensatory postsaccadic ocular drift. This ocular drift was still evident in the dark, although smaller (typically 15% of the amplitude of the antecedent saccade, up to a maximum drift of 8 degrees). Retinal slip alone, without a net displacement of the image, was sufficient to elicit these adaptive changes, and compensation for leftward and rightward saccades was independent. It took several days to complete adaptation, but recovery (in the light) was much quicker. The decay of this adaptation in darkness was very slow; after 3 days the ocular drift was reduced by less than 50%. The time constants of single exponential curve fits to adaptation time courses of data from five animals were 35 h for acquisition, 4 h for recovery, and at least 40 h for decay in darkness. Descriptions of the central innervation for a saccade are usually simplified to only two components: a pulse and a step. It has been hypothesized that suppression of pathological postsaccadic drift is achieved by adjusting the ratio of the pulse to the step of innervation (19, 26). However, we show that the time constant of the ocular drift is influenced by the time constant of the adapting stimulus, which cannot be explained by the simple pulse-step model of saccadic innervation. A more realistic representation of the saccadic innervation has three components: a pulse, an exponential slide, and a step. Normal saccades were accurately simulated by a fourth-order, linear model of the ocular motor plant driven by such a pulse-slide-step combination. Saccades made after prolonged exposure to optically induced retinal image slip could also be simulated by properly adjusting the slide and step components.(ABSTRACT TRUNCATED AT 400 WORDS)

Adaptation, Ocular↗