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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
PubMed2006

Human vergence eye movements initiated by competing disparities: evidence for a winner-take-all mechanism.

Vergence eye movements were elicited in human subjects at short latencies (approximately 70 ms) by applying binocular disparities briefly (200 ms) to large grating patterns (46 degrees wide, 35 degrees high). The positions of both eyes were recorded with the electromagnetic search coil technique. Using a dichoptic viewing arrangement (Wheatstone stereoscope), each eye viewed two overlapping 1-D sine waves that had the same orientation but different spatial frequencies. These two sine waves each had a binocular disparity that was 1/4 of its wavelength and the effect of varying their relative contrasts was examined (15 contrast ratios ranging from 0.125 to 8). The first experiment used horizontal gratings and recorded the vertical vergence responses when the two sine waves had spatial frequencies in the ratio 3:5 and vertical disparities of opposite sign. Initial vergence responses showed a highly nonlinear dependence on the contrast ratio. On average, when the contrast of one sine wave exceeded that of the other by a factor of >2.2, the sine wave with the higher contrast dominated responses and the sine wave with the lower contrast had almost no influence: winner-take-all. A second experiment, which used vertical gratings and recorded the horizontal vergence responses when the two sine waves had spatial frequencies in the ratio 3:5 and horizontal disparities of opposite sign, also uncovered nonlinear interactions but these were much more variable from one subject to another and, on average, one sine wave did not achieve complete dominance until its contrast exceeded that of the other by a factor of >4.5. When these two experiments were repeated with grating patterns in which the two sine waves had spatial frequencies in the ratio 3:7 and disparities of the same sign, similar nonlinear interactions were apparent. We attribute the nonlinear dependence on relative contrast to mutual inhibition between the neural elements processing the disparities of the two sine waves. We further suggest that this interaction will help to maintain binocular alignment on the objects in the plane of regard because the retinal images of those objects will tend to be better focused-and hence tend to have higher contrasts-than the images of objects in other depth planes.

Contrast Sensitivity↗
PubMed2006

Short-latency disparity vergence eye movements: a response to disparity energy.

Vergence eye movements were elicited in human subjects by applying disparities to square-wave gratings lacking the fundamental ("missing fundamental", mf). Using a dichoptic arrangement, subjects viewed gratings that were identical at the two eyes except for a phase difference of 1/4 wavelength so that, based on the nearest-neighbor matches, the features and the 4n+1 harmonics (5th, 9th, etc.) all had binocular disparities of one sign, whereas the 4n-1 harmonics (3rd, 7th, etc.) all had disparities of the opposite sign. Further, the amplitude of the ith harmonic was proportional to 1/i. Using the electromagnetic search coil technique to record the positions of both eyes indicated that the earliest vergence eye movements elicited by these disparity stimuli had ultra-short latencies (minimum, <65 ms) and were always in the direction of the most prominent harmonic, the 3rd, but their magnitudes fell short of those elicited when the same disparities were applied to pure sinusoids whose spatial frequency and contrast matched those of the 3rd harmonic. This shortfall was evident in both the horizontal vergence responses recorded with vertical grating stimuli and the vertical vergence responses recorded with horizontal grating stimuli. When the next most prominent harmonic, the 5th, was removed from the mf stimulus (creating the "mf-5" stimulus) the vertical vergence responses showed almost no shortfall-indicating that it had been almost entirely due to that 5th harmonic-but the horizontal vergence responses still showed a small shortfall, at least with higher contrast stimuli. This small shortfall might represent a very minor contribution from higher harmonics and/or distortion products and/or a feature-based mechanism. We conclude that the earliest disparity vergence responses-especially vertical-were strongly dependent on the major Fourier components of the binocular images, consistent with early spatial filtering of the monocular visual inputs prior to their binocular combination as in the disparity-energy model of complex cells in striate cortex [Ohzawa, I., DeAngelis, G. C., & Freeman, R. D. (1990). Stereoscopic depth discrimination in the visual cortex: neurons ideally suited as disparity detectors. Science, 249, 1037-1041].

Computer Graphics↗
PubMed2006

Human ocular following initiated by competing image motions: evidence for a winner-take-all mechanism.

The initial ocular following responses (OFRs) elicited by 1/4-wavelength steps applied to the missing fundamental (mf) stimulus are in the backward direction and largely determined by the principal Fourier component, the 3rd harmonic [Sheliga, B. M., Chen, K. J., FitzGibbon, E. J., & Miles, F. A. (2005). Initial ocular following in humans: A response to first-order motion energy. Vision Research, 45, 3307-3321]. When the contrast of the 3rd harmonic was selectively reduced below that of the next most prominent harmonic-the 5th, which moves in the opposite (forward) direction-then the OFR reversed direction and the 3rd harmonic effectively lost all of its influence as the OFR was now largely determined by the 5th harmonic. Restricting the stimulus to just two sine waves (of equal efficacy when of equal contrast and presented singly) with the spatial frequencies of the 3rd and 5th harmonics of the mf stimulus indicated that the critical factor was the ratio of their two contrasts: when of similar contrast both were effective (vector sum/averaging), but when the contrast of one was <1/2 that of the other then the one with the lower contrast became ineffective (winner-take-all). This nonlinear dependence on the contrast ratio was attributed to mutual inhibition and was well described by a weighted-average model with just two free parameters. Further experiments with broadband and dual-grating stimuli indicated that nonlinear interactions occur not only in the neural processing of stimuli moving in opposite directions but also of stimuli that share the same direction and differ only in their spatial frequency and speed. Clearly, broad-band and dual-grating stimuli can uncover significant nonlinearities in visual information processing that are not evident with single sine-wave stimuli.

Contrast Sensitivity↗
PubMed2005

The initial ocular following responses elicited by apparent-motion stimuli: reversal by inter-stimulus intervals.

Transient apparent-motion stimuli, consisting of single 1/4-wavelength steps applied to square-wave gratings lacking the fundamental ("missing fundamental stimulus") and to sinusoidal gratings, were used to elicit ocular following responses (OFRs) in humans. As previously reported [Sheliga, B. M., Chen, K. J., FitzGibbon, E. J., & Miles, F. A. (2005). Initial ocular following in humans: a response to first-order motion energy. Vision Research, in press], the earliest OFRs were strongly dependent on the motion of the major Fourier component, consistent with early spatio-temporal filtering prior to motion detection, as in the well-known energy model of motion analysis. Introducing inter-stimulus intervals (ISIs) of 10-200 ms, during which the screen was gray with the same mean luminance, reversed the initial direction of the OFR, the peak reversed responses (with ISIs of 20-40 ms) being substantially greater than the non-reversed responses (with an ISI of 0 ms). When the mean luminance was reduced to scotopic levels, reversals now occurred only with ISIs > or=60 ms and the peak reversed responses (with ISIs of 60-100 ms) were substantially smaller than the non-reversed responses (with an ISI of 0 ms). These findings are consistent with the idea that initial OFRs are mediated by first-order motion-energy-sensing mechanisms that receive a visual input whose temporal impulse response function is strongly biphasic in photopic conditions and almost monophasic in scotopic conditions.

Contrast Sensitivity↗
PubMed2005

Initial ocular following in humans: a response to first-order motion energy.

Visual motion is sensed by low-level (energy-based) and high-level (feature-based) mechanisms. Ocular following responses (OFR) were elicited in humans by applying horizontal motion to vertical square-wave gratings lacking the fundamental ("missing fundamental stimulus"). Motion consisted of successive 1/4-wavelength steps, so the features and 4n+1 harmonics (where n=integer) shifted forwards, whereas the 4n-1 harmonics--including the strongest Fourier component (the 3rd harmonic)--shifted backwards (spatial aliasing). Initial OFR, recorded with the electromagnetic search coil technique, were always in the direction of the 3rd harmonic, e.g., leftward steps resulted in rightward OFR. Thus, the earliest OFR were strongly dependent on the motion of the major Fourier component, consistent with early spatio-temporal filtering prior to motion detection, as in the well-known energy model of motion analysis.

Contrast Sensitivity↗
PubMed2005

Short-latency disparity vergence in humans: evidence for early spatial filtering.

Our study was concerned with the disparity detectors underlying the initial disparity vergence responses (DVRs) that are elicited at ultrashort latencies by binocular disparities applied to large images. DVRs were elicited in humans by applying horizontal disparity to vertical square-wave gratings lacking the fundamental (termed here, the "missing fundamental"). In the frequency domain, a pure square wave is composed of odd harmonics--first, third, fifth, seventh, etc.--such that the third, fifth, seventh, etc., have amplitudes that are one-third, one-fifth, one-seventh, etc., that of the first, and the missing fundamental lacks the first harmonic. The patterns seen by the two eyes have a phase difference of one-quarter wavelength, so the disparity of the features and 4n + 1 harmonics (where n = integer) has one sign (crossed or uncrossed), whereas the 4n - 1 harmonics--including the strongest Fourier component (the third harmonic)--has the opposite sign (uncrossed or crossed): spatial aliasing. The earliest DVRs, recorded with the search-coil technique, had minimum latencies of 70 to 80 ms and were generally in the direction of the third harmonic, that is, uncrossed disparities resulted in convergent eye movements. In other experiments on the DVRs, one eye saw a missing fundamental and the other saw a pure sine wave with the contrast and wavelength of the third harmonic but differing in phase by one-quarter wavelength. This resulted in short-latency vergence in accordance with matching of the third harmonic. These data all indicate the importance of the Fourier components, consistent with early spatial filtering prior to binocular matching.

Fourier Analysis↗
PubMed2005

Initial ocular following in humans depends critically on the fourier components of the motion stimulus.

Visual motion is sensed by low-level (energy-based) and high-level (feature-based) mechanisms. Our interest is in the motion detectors underlying the initial ocular following responses (OFR) that are elicited at ultrashort latencies by sudden motions of large images. OFR were elicited in humans by applying horizontal motion to vertical square-wave gratings lacking the fundamental. In the frequency domain, a pure square wave is composed of the odd harmonics--first, third, fifth, seventh, etc.--such that the third, fifth, seventh, etc., have amplitudes that are one-third, one-fifth, one-seventh, etc., that of the first, and the missing fundamental stimulus lacks the first harmonic. Motion consisted of successive quarter-wavelength steps, so the features and 4n+1 harmonics (where n = integer) shifted forward, whereas the 4n-1 harmonics--including the strongest Fourier component (the third harmonic)--shifted backward (spatial aliasing). Thus, the net Fourier energy and the non-Fourier features moved in opposite directions. Initial OFR, recorded with the search coil technique, had minimum latencies of 60 to 70 ms and were always in the direction of the third harmonic, for example, leftward steps resulted in rightward OFR. Thus, the earliest OFR were strongly dependent on the motion of the major Fourier component, consistent with mediation by oriented spatiotemporal visual filters as in the well-known energy model of motion detection. Introducing interstimulus intervals of 10 to 100 ms (during which the screen was uniform gray) reversed the initial direction of tracking, consistent with extensive neurophysiological and psychophysical data suggesting that the visual input to the motion detectors has a biphasic temporal impulse response.

Fourier Analysis↗
PubMed2003

Short-latency disparity-vergence eye movements in humans: sensitivity to simulated orthogonal tropias.

Small disparity stimuli applied to large random-dot patterns elicit machine-like vergence eye movements at short latency. We have examined the sensitivity of these eye movements to simulated orthogonal tropias in three normal subjects by recording (1) the effects of vertical disparities on the initial horizontal vergence responses elicited by 2 degrees crossed and uncrossed (horizontal) disparity stimuli, and (2) the effects of horizontal disparities on the initial vertical vergence responses elicited by 1.2 degrees left-hyper and 0.8 degrees right-hyper (vertical) disparity stimuli. Initial vergence responses were strongest when the orthogonal disparity was close to zero, and decreased to zero as the orthogonal disparity increased to 3 degrees -5 degrees, i.e., there was only a limited tolerance for orthogonal disparity. Tuning curves describing the dependence of the initial change in the vergence angle on the orthogonal disparity were well fit by a Gaussian function. An additional subject, who had an esotropia of approximately 10 degrees in our experimental setup, showed almost no horizontal vergence responses but did show vertical vergence responses to vertical disparity stimuli at short latency (albeit slightly longer than normal) despite the fact that her esotropia resulted in uncrossed disparities that would have totally disabled the vertical vergence mechanism of a normal subject, cf., anomalous retinal correspondence.

Convergence, Ocular↗
PubMed2003

Short-latency ocular following in humans is dependent on absolute (rather than relative) binocular disparity.

A previous study showed that the initial ocular following responses elicited by sudden motion of a large random-dot pattern were only modestly attenuated when that whole pattern was shifted out of the plane of fixation by altering its horizontal binocular disparity, but the same disparity applied to a restricted region of the dots had a much more powerful effect [Vision Research 41 (2001) 3371]. Thus, if the dots were partitioned into horizontal bands, for example, and alternate bands were moved in opposite directions to the left or right then ocular following was very weak, but if the (conditioning) dots moving in one direction were all shifted out of the plane of fixation (by applying horizontal disparity to them) then strong ocular following was now seen in the direction of motion of the (test) dots in the plane of fixation, i.e., moving images became much less effective when they were given binocular disparity. We sought to determine if the greater impact of disparity with the partitioned images was because there were additional relative disparity cues. We used a similar partitioned display and found that the dependence of ocular following on the absolute disparity of the conditioning stimulus had a Gaussian form with an x-offset that was close to zero disparity and, importantly, this offset was almost unaffected by changing the absolute disparity of the test stimulus. We conclude from this that it is the absolute--rather than the relative--disparity that is important, and that ocular following has a strong preference for moving images whose absolute disparities are close to zero. This is consistent with the idea that ocular following selectively stabilizes the retinal images of objects in and around the plane of fixation and works in harmony with disparity vergence, which uses absolute disparity to bring objects of interest into the plane of fixation [Archives of Ophthalmology 55 (1956) 848].

Eye Movements↗
PubMed2002

Reversed short-latency ocular following.

Using the scleral search coil technique to monitor eye movements, we recorded short-latency ocular following responses to displacement steps of large random-dot patterns. On half of the trials, the luminance of the dots and background were reversed during the step, a procedure that is known to reverse the direction of the perceived motion ("reverse phi"). Steps without luminance reversal induced small but consistent ocular following in the direction of the steps at ultra-short latency (<80 ms). Steps with luminance reversal induced small but consistent tracking at the same latency but in the direction opposite to the actual displacement. Tuning curves describing the dependence of initial ocular following on the amplitude of the displacement had a form approximating the derivative of a Gaussian and were well fit by Gabor functions, the cosine term being phase shifted approximately 180 degrees by the luminance reversal. This result is consistent with the idea that the initial ocular following is mediated, at least in part, by first-order (luminance) motion-energy detectors.

Eye Movements↗
PubMed2002

Voluntary saccadic eye movements in humans studied with a double-cue paradigm.

In the classic double-step paradigm, subjects are required to make a saccade to a visual target that is briefly presented at one location and then shifted to a new location before the subject has responded. The saccades in this situation are "reflexive" in that they are made in response to the appearance of the target itself. In the present experiments we adapted the double-step paradigm to study "voluntary" saccades. For this, several identical targets were always visible and subjects were given a cue to indicate that they should make a saccade to one of them. This cue was then changed to indicate another of the targets before the subject had responded: double-cue (DC) paradigm. The saccadic eye movements in our DC paradigm had many features in common with those in the double-step paradigm and we show that apparent differences can be attributed to the spatio-temporal arrangements of the cues/targets rather than to any intrinsic differences in the programming of these two kinds of eye movements. For example, a feature of our DC paradigm that is not seen in the usual double-step paradigm is that the second cue could cause transient delays of the initial saccade, and these delays still occurred when the second cue was reflexive--provided that it was at the fovea (as in our DC paradigm) and not in the periphery (as in the usual double-step paradigm). Thus, the critical factor for the delay was the retinal (foveal) location of the second cue/target--not whether it was cognitive or reflexive--and we argue that the second cue/target is here acting as a distractor. We conclude that the DC paradigm can be used to study the programming of voluntary saccades in the same way that the double-step paradigm can be used to study reflexive saccades.

Cognition↗
PubMed2002

Version and vergence eye movements in humans: open-loop dynamics determined by monocular rather than binocular image speed.

We examined the velocity dependence of the vergence and version eye movements elicited by motion stimuli that were symmetric or asymmetric at the two eyes. Movements of both eyes were recorded with the scleral search coil technique. Vergence was computed as the difference in the positions of the two eyes (left-right) and version was computed as the average position of the two eyes ((left+right)/2). Subjects faced a large tangent screen onto which two identical random-dot patterns were back-projected. Each pattern was viewed by one eye only using crossed-polarizers and its position was controlled by X/Y mirror galvanometers. Viewing was always binocular and horizontal velocity steps (range, 5-240 deg/s) were applied to one (asymmetric stimulus) or both (symmetric stimulus) patterns approximately 50 ms after a centering saccade. With the symmetric stimulus, the motion at the two eyes could be either in the opposite direction (eliciting vergence responses) or in the same direction (eliciting version responses). The asymmetric stimuli elicited both vergence and version. In all cases, minimum response latencies were very short (<90 ms). Velocity tuning curves (based on the changes in vergence and version over the time period, 90-140 ms) were all sigmoidal and peaked when the monocular (i.e., retinal) image velocities were 30-60 deg/s. The vergence (version) responses to symmetric stimuli were linearly related to the vergence (version) responses to asymmetric stimuli when expressed in terms of the monocular rather than the binocular image velocities. We conclude that the dynamical limits for both vergence and version are imposed in the monocular visual pathways, before the inputs from the two eyes are combined.

Eye Movements↗
PubMed2002

Population coding in cortical area MST.

Disparity steps applied to large patterns elicit vergence eye movements at ultrashort latencies. Disparity tuning curves, describing the dependence of the amplitude of the initial vergence responses on the amplitude of the disparity steps, resemble the derivative of a gaussian and indicate that appropriate servo-like behavior occurs only with small disparity steps (<1 degree). Lesion data from monkeys suggest that these vergence responses are mediated, at least in part, by neurons in the medial superior temporal area of the cerebral cortex, and we here review a recent study of the associated single unit activity in that area. Few medial superior temporal neurons have disparity tuning curves whose shapes resemble the tuning curve for vergence. Yet, when the disparity tuning curves for all of the disparity-sensitive cells recorded from a given monkey are summed together, they match the tuning curves for the vergence responses of that monkey very closely, even reproducing that animal's idiosyncracies. When all of the spike trains elicited by a given disparity step are summed together to give an average discharge profile for the whole population of recorded cells, many are noisy, but others that are less so match the temporal profile of the motor response, vergence velocity, quite well. We conclude that the discharges of the disparity-sensitive cells in the medial superior temporal area each represent only a very limited aspect of the sensory stimulus (and/or associated motor response?), but when pooled together, they provide a complete description of the vergence velocity motor response: population coding.

Animals↗
PubMed2001

Short-latency ocular following in humans: sensitivity to binocular disparity.

We show that the initial ocular following responses elicited by motion of a large pattern are modestly attenuated when that pattern is shifted out of the plane of fixation by altering its binocular disparity. If the motion is applied to only restricted regions of the pattern, however, then altering the disparity of those regions severely attenuates their ability to generate ocular following. This sensitivity of the ocular tracking mechanism to local binocular disparity would help the observer who moves through a cluttered 3-D world to stabilize objects in the plane of fixation and ignore all others.

Eye Movements↗
PubMed2001

Short-latency disparity vergence in humans.

Eye movement recordings from humans indicated that brief exposures (200 ms) to horizontal disparity steps applied to large random-dot patterns elicit horizontal vergence at short latencies (80.9 +/- 3.9 ms, mean +/- SD; n = 7). Disparity tuning curves, describing the dependence of the initial vergence responses (measured over the period 90-157 ms after the step) on the magnitude of the steps, resembled the derivative of a Gaussian, with nonzero asymptotes and a roughly linear servo region that extended only a degree or two on either side of zero disparity. Responses showed transient postsaccadic enhancement: disparity steps applied in the immediate wake of saccadic eye movements yielded higher vergence accelerations than did the same steps applied some time later (mean time constant of the decay, 200 ms). This enhancement seemed to be dependent, at least in part, on the visual reafference associated with the prior saccade because similar enhancement was observed when the disparity steps were applied in the wake of saccadelike shifts of the textured pattern. Vertical vergence responses to vertical disparity steps were qualitatively similar: latencies were longer (on average, by 3 ms), disparity tuning curves had the same general form but were narrower (by approximately 20%), and their peak-to-peak amplitudes were smaller (by approximately 70%). Initial vergence responses usually had directional errors (orthogonal components) with a very systematic dependence on step size that often approximated an exponential decay to a nonzero asymptote (mean space constant +/- SD, 1.18 +/- 0.66 degrees ). Based on the asymptotes of these orthogonal responses, horizontal errors (with vertical steps) were on average more than three times greater than vertical errors (with horizontal steps). Disparity steps >7 degrees generated "default" responses that were independent of the direction of the step, idiosyncratic, and generally had both horizontal and vertical components. We suggest that the responses depend on detectors that sense local disparity matches, and that orthogonal and "default" responses result from globally "false" matches. Recordings from three monkeys, using identical disparity stimuli, confirmed that monkeys also show short-latency disparity vergence responses (latency approximately 25 ms shorter than that of humans), and further indicated that these responses show all of the major features seen in humans, the differences between the two species being solely quantitative. Based on these data and those of others implying that foveal images normally take precedence, we suggest that the mechanisms under study here ordinarily serve to correct small vergence errors, automatically, especially after saccades.

Adult↗
PubMed2001

Single-unit activity in cortical area MST associated with disparity-vergence eye movements: evidence for population coding.

Single-unit discharges were recorded in the medial superior temporal area (MST) of five behaving monkeys. Brief (230-ms) horizontal disparity steps were applied to large correlated or anticorrelated random-dot patterns (in which the dots had the same or opposite contrast, respectively, at the two eyes), eliciting vergence eye movements at short latencies [65.8 +/- 4.5 (SD) ms]. Disparity tuning curves, describing the dependence of the initial vergence responses (measured over the period 50-110 ms after the step) on the magnitude of the steps, resembled the derivative of a Gaussian, the curves obtained with correlated and anticorrelated patterns having opposite sign. Cells with disparity-related activity were isolated using correlated stimuli, and disparity tuning curves describing the dependence of these initial neuronal responses (measured over the period of 40-100 ms) on the magnitude of the disparity step were constructed (n = 102 cells). Using objective criteria and the fuzzy c-means clustering algorithm, disparity tuning curves were sorted into four groups based on their shapes. A post hoc comparison indicated that these four groups had features in common with four of the classes of disparity-selective neurons in striate cortex, but three of the four groups appeared to be part of a continuum. Most of the data were obtained from two monkeys, and when the disparity tuning curves of all the individual neurons recorded from either monkey were summed together, they fitted the disparity tuning curve for that same animal's vergence responses remarkably well (r(2): 0.93, 0.98). Fifty-six of the neurons recorded from these two monkeys were also tested with anticorrelated patterns, and all showed significant modulation of their activity (P < 0.005, 1-way ANOVA). Further, when all of the disparity tuning curves obtained with these patterns from either monkey were summed together, they too fitted the disparity tuning curve for that same animal's vergence responses very well (r(2): 0.95, 0.96). Indeed, the summed activity even reproduced idiosyncratic differences in the vergence responses of the two monkeys. Based on these and other observations on the temporal coding of events, we hypothesize that the magnitude, direction, and time course of the initial vergence velocity responses associated with disparity steps applied to large patterns are all encoded in the summed activity of the disparity-sensitive cells in MST. Latency data suggest that this activity in MST occurs early enough to play an active role in the generation of vergence eye movements at short latencies.

Algorithms↗
PubMed1999

Short-latency visual stabilization mechanisms that help to compensate for translational disturbances of gaze.

Recent studies in primates have revealed short-latency visual tracking mechanisms that help to stabilize the eyes during translational disturbances of the observer, and so operate as backups to otolith-mediated vestibulo-ocular reflexes. One such mechanism generates version eye movements to help stabilize gaze when the moving observer looks off to one side, utilizing binocular disparity to help single out the images in the plane of fixation (ocular following). Two others generate vergence eye movements to help maintain binocular alignment on objects that lie ahead: one responds to the radial patterns of optic flow (radial-flow vergence) and the other to the changes in binocular parallax (disparity vergence). Accumulating evidence suggests that, despite their short latency, all are mediated by the medial superior temporal area of cortex.

Adaptation, Physiological↗