Scott B. Stevenson

Discrimination of jittered sonar echoes by the echolocating bat, Eptesicus fuscus: the shape of target images in echolocation.

Summary1.Behavioral experiments with jittering echoes examined acoustic images of sonar targets in the echolocating bat, Eptesicus fuscus, along the echo delay or target range axis. Echo phase, amplitude, bandwidth, and signal-to-noise ratio were manipulated to assess the underlying auditory processes for image formation.2.Fine delay acuity is about 10 ns. Calibration and control procedures indicate that this represents temporal acuity rather than spectral discrimination. Jitter discrimination curves change in phase when the phase of one jittering echo is shifted by 180° relative to the other, showing that echo phase is involved in delay estimation. At an echo detectability index of about 36 dB, fine acuity is 40 ns, which is approximately as predicted for the delay accuracy of an ideal receiver.3.Compound performance curves for 0° and 180° phase conditions match the crosscorrelation function of the echoes. The locations of both 0° and 180° phase peaks in the performance curves shift along the time axis by an amount that matches neural amplitude-latency trading in Eptesicus, confirming a temporal basis for jitter discrimination.

Interocular correlation, luminance contrast and cyclopean processing

We have investigated the nature and viability of interocular correlation as a measure of signal strength in the cyclopean domain. Thresholds for the detection of interocular correlation in dynamic random element stereograms were measured as a function of luminance contrast, a more traditional measure of stimulus strength. At high contrasts, correlation thresholds were independent of contrast. At low contrasts, correlation thresholds were inversely proportional to the square of contrast. Stereothresholds were also measured as a function of both contrast and interocular correlation. At low contrasts, stereoacuity was inversely proportional to both interocular correlation and the square of contrast. These results are consistent with an inherently multiplicative mechanism of binocular combination, such as a cross-correlation of the two eyes inputs.

Correcting for miniature eye movements in high resolution scanning laser ophthalmoscopy

The newest generation of confocal scanning laser ophthalmoscopes with adaptive optics correction of ocular aberrations provides retinal images of unprecedented resolution, allowing for real-time imaging of photoreceptors in the living human eye. Natural fixational eye movements made by the subject/patient during recording produce distortions that are unique in each frame. Correction for these distortions is necessary before multiple frames can be added together to achieve noise reduction or to build a mosaic image from different retinal areas. Here we describe the characteristics of fixational eye movements and the distortions they produce during retinal imaging, we show examples of images with particular distortions, and show eye movement records obtained during the correction of these distortions.

Negative feedback control model of proximal convergence and accommodation

A comprehensive model has been developed to illustrate the interactions between the observer and the surrounding environment in the control of oculomotor responses to distance or 3‐D space. Accommodation and vergence respond to both spatiotopic (body referenced) proximal percepts and retinotopic (eye referenced) physical stimuli of blur and disparity. Both spatiotopic and retinotopic stimuli are derived respectively from perceptual and physical correlates of negative feedback for eye position. The spatiotopic and retinotopic stimulus errors are combined in the feed forward path and drive a common occulomotor controller which has a phasic‐tonic organization. Spatiotopic and retinotopic stimuli are shown to be effective over complementary operating ranges. Perceptual spatiotopic errors of gaze provide optimal stimuli for near responses to large depth intervals whereas physical‐retinotopic cues of blur and disparity provide quantitative information about small binocular fixation errors. Small dynamic variations of target distance are sensed both spatiotopically and retinotopically. Coarse and fine spatiotopic errors of gaze are processed differently. Large spatiotopic errors are sampled intermittently al the beginning of the near response, whereas small retinotopic position errors and spatiotopic velocity errors are sampled continuously throughout the near response. Former reports of empirically observed higher velocity of vergence responses to very large depth intervals is explained in terms of stimulus sampling modes rather than in terms of separate oculomotor control mechanisms. The model demonstrates a complementary function of top‐down spatiotopic cues, which are used to initiate the near response, and bottom‐up retinotopic cues, which are used to refine and complete the near response. Cross‐couplings by vergence‐accommodation and accommodative‐vergence serve to coordinate the components of the near response when feedback from the sensed response of one motor system (i.e. vergence) is more accurate than that of the other motor system (i.e. accommodation). The model presented here is concerned primarily with the near response mediated by accommodation and disjunctive eye movements and not by the independent vergence mediated by non‐conjugate or yoked saccades of unequal amplitude.

Disparity tuning in mechanisms of human stereopsis

The change in sensitivity across some stimulus dimension which follows adaptation to a particular stimulus can reveal a great deal about the tuning characteristics of underlying sensory/perceptual mechanisms. In this study, a psychophysical adaptation paradigm was employed to characterize the disparity tuning of perceptual mechanisms involved in stereopsis. The stimulus was a dynamic random-dot stereogram (DRDS) portraying a surface which varied in interocular correlation (IOC) and retinal disparity. Adaptation to a fully correlated DRDS surface produced an elevation in IOC threshold over a relatively narrow range of disparities, with maximum effect at the disparity of the adapting stimulus. The width of these disparity tuning functions varied from 5 arc min for adaptation at the horopter to 20 arc min for adaptation at 20 arc min disparity. Frequently, IOC sensitivity was enhanced for disparities on either side of the adapted disparity, suggesting that an opponent center-surround organization operates at an early level of disparity processing. A model of underlying channel structure consistent with these data is presented.

Clutter interference and the integration time of echoes in the echolocating bat, Eptesicus fuscus

The ability of the echolocating bat, Eptesicus fuscus, to detect a sonar target is affected by the presence of other targets along the same axis at slightly different ranges. If echoes from one target arrive at about the same delay as echoes from another target, clutter interference occurs and one set of echoes masks the other. Although the bats sonar emissions and the echoes themselves are 2 to 5 ms long, echoes (of approximately equal sensation levels--around 15 dB SL) only interfere with each other if they arrive within 200 to 400 microseconds of the same arrival time. This figure is an estimate of the integration time of the bats sonar receiver for echoes. The fine structure of the clutter-interference data reflects the reinforcement and cancellation of echoes according to their time separation. When clutter interference first occurs, the waveforms of test and cluttering echoes already overlap for much of their duration. The masking effect underlying clutter interference appears specifically due to overlap, not between raw echo waveforms, but between the patterns of mechanical excitation created when echoes pass through bandpass filters equivalent to auditory-nerve tuning curves. While the time scale of clutter interference is substantially shorter than the duration of echo waveforms, it still is much longer than the eventual width of a targets range-axis image expressed in terms of echo delay.

Isovergence surfaces: the conjugacy of vertical eye movements in tertiary positions of gaze

Conjugate gaze is often defined as the equal angle rotation of the two eyes. For fixation at far distances, the optical axes are parallel and conjugacy is defined irrespective of the coordinate system. For nearby or finite fixation distances, the evaluation of conjugacy for many gaze postures depends on the coordinate system used to measure it. For example, if the eye is elevated or depressed and the eye is rotated about a vertical axis, the intersections of lines of sight with a tangent screen will describe either straight lines for arcs depending on whether the vertical axis is fixed with respect to the head or to the eye. Because of the horizontal separation of the two eyes, the binocular fixation of near targets at tertiary positions of gaze will require a vertical vergence component for head‐referenced but not eye‐referenced measurements. The vertical gaze alignment of three human subjects was measured as they viewed targets placed at secondary and tertiary eye positions at two different distances. Vertical vergence was either held open or closed‐loop. The lines of sight were found lo intersect (i.e. vertical gaze was aligned) regardless of target position or viewing condition.

Hyperacuity, superresolution and gap resolution in human stereopsis

Different types of stereoscopic acuity were studied with tasks adapted from studies of visual direction acuity. Dynamic, random-element stereograms portraying multiple surfaces in depth and a temporal 2AFC procedure were used for all measurements. The three tasks required detection of a depth offset (Hyperacuity task), a depth-axis thickening (Superresolution task), and a depth-axis gap between surfaces (Gap Resolution task). Thresholds for the three tasks were on the order of 3 sec arc, 30 sec arc and 200 sec arc of retinal disparity, respectively. These results are comparable to those for the analogous visual direction tasks on which they were patterned, suggesting that the underlying judgments involved are similar. Results are used to estimate the intrinsic noise of horizontal disparity processing.

Depth attraction and repulsion in random dot stereograms

Previous studies of perceived attraction or repulsion of adjacent visual targets have used local targets whose positions were varied in both depth and direction. We have measured these effects in three subjects using dynamic random-dot stereograms to isolate depth-axis effects. Results show that both attraction and repulsion effects can occur for overlapping, positively correlated, random-dot surfaces. The results were quantitatively similar to those reported previously for local targets. Manipulation of interocular correlation confirmed that the effects are produced by binocular interactions. Results are explained as accurate judgments based on the stimulus at the cyclopean level.

The Influence of Subject Instruction on Horizontal and Vertical Vergence Tracking

Previously it has been reported that horizontal disparity vergence is strongly influenced by subject instructions to vary attention or tracking effort. This paper describes experiments which compared these instruction effects on horizontal and vertical disparity vergence. Within-trial comparisons were made possible by use of oblique (combined horizontal and vertical) disparity modulation. Subjects viewed a flat, fully correlated, dynamic random noise stereogram pattern through stationary circular apertures, with a small stationary fixation cross superimposed in the center. The disparity of the noise pattern was either modulated sinusoidally or changed abruptly. Subjects were instructed either to (1) hold fixation on the cross and ignore the disparity modulation of the noise pattern; or (2) follow the movement of the noise pattern as accurately as possible. Subjects showed clear effects of instruction on the horizontal component of tracking, but showed little or no effect on the vertical component. Horizontal and vertical components of oblique vergence tracking appear to be largely independent, and vertical vergence is affected minimally, if at all, by an effort to track.