Seeing the future: predictive control in neural models of ocular accommodation

Ocular accommodation is the process of adjusting the eye's crystalline lens so as to bring the retinal image into sharp focus. The major stimulus to accommodation is therefore retinal defocus, and in essence, the job of accommodative control is to send a signal to the ciliary muscle which will minimise the magnitude of defocus. In this paper, we first provide a tutorial introduction to control theory to aid vision scientists without this background. We then present a unified model of accommodative control that explains properties of the accommodative response for a wide range of accommodative stimuli. Following previous work, we conclude that most aspects of accommodation are well explained by dual integral control, with a "fast" or "phasic" integrator enabling response to rapid changes in demand, which hands over control to a "slow" or "tonic" integrator which maintains the response to steady demand. Control is complicated by the sensorimotor latencies within the system, which delay both information about defocus and the accommodation changes made in response, and by the sluggish response of the motor plant. These can be overcome by incorporating a Smith predictor, whereby the system predicts the delayed sensory consequences of its own motor actions. For the first time, we show that critically-damped dual integral control with a Smith predictor accounts for adaptation effects as well as for the gain and phase for sinusoidal oscillations in demand. In addition, we propose a novel proportional-control signal to account for the power spectrum of accommodative microfluctuations during steady fixation, which may be important in hunting for optimal focus, and for the nonlinear resonance observed for low-amplitude, high-frequency input. Complete Matlab/Simulink code implementing the model is provided at https://doi.org/10.25405/data.ncl.14945550

Impact of microsaccades on visual shape processing

Sensitivity to subtle changes in the shape of visual objects has been attributed to the existence of global pooling mechanisms that integrate local form information across space. While global pooling is typically demonstrated under steady fixation, other work suggests prolonged fixation can lead to a collapse of global structure. Here we ask whether small ballistic eye movements that naturally occur during periods of fixation affect the global processing of radial frequency (RF) patterns - closed contours created by sinusoidally modulating the radius of a circle. Observers were asked to discriminate the shapes of circular and RF modulated patterns while fixational eye movements were recorded binocularly at 500Hz. Microsaccades were detected using a velocity-based algorithm, allowing trials to be sorted according to the relative timing of stimulus and microsaccade onset. Results revealed clear peri-saccadic changes in shape discrimination thresholds. Performance was impaired when microsaccades occurred close to stimulus onset, but facilitated when they occurred shortly afterwards. In contrast, global integration of shape was unaffected by the timing of microsaccades. These findings suggest that microsaccades alter the discrimination sensitivity to briefly presented shapes but do not disrupt the spatial pooling of local form signals.

The Rotating Snakes Illusion Is a Straightforward Consequence of Nonlinearity in Arrays of Standard Motion Detectors

The Rotating Snakes illusion is a motion illusion based on repeating, asymmetric luminance patterns. Recently, we found certain gray-value conditions where a weak illusory motion occurs in the opposite direction. Of the four models for explaining the illusion, one also explains the unexpected perceived opposite direction.We here present a simple new model, without free parameters, based on an array of standard correlation-type motion detectors with a subsequent nonlinearity (e.g., saturation) before summing the detector outputs. The model predicts (a) the pattern-appearance motion illusion for steady fixation, (b) an illusion under the real-world situation of saccades across or near the pattern (pattern shift), (c) a relative maximum of illusory motion for the same gray values where it is found psychophysically, and (d) the opposite illusion for certain luminance values. We submit that the new model’s sparseness of assumptions justifies adding a fifth model to explain this illusion.

The Rotating Snake Illusion is a straightforward consequence of non-linearity in arrays of standard motion detectors

The Rotating Snakes illusion is a motion illusion based on repeating, asymmetric luminance patterns. Recently, we found certain grey-value conditions where a weak, illusory motion occurs in the opposite direction. Of the four models for explaining the illusion, one (Backus and Oruç, 2005) also explains the unexpected perceived opposite direction. We here present a simple new model, without free parameters, based on an array of standard correlation-type motion detectors with a subsequent non-linearity (e.g., saturation) before summing the detector outputs. The model predicts (1) the pattern-appearance motion illusion for steady fixation, (2) an illusion under the real-world situation of saccades across or near the pattern (pattern shift), (3) a relative maximum of illusory motion for the same grey values where it is found psychophysically, and (4) the inverse illusion for certain luminance values. We submit that the model’s sparseness of assumptions justifies adding a fifth model to explain this illusion.

Saccadic Suppression of Displacement Does Not Reflect a Saccade-Specific Bias to Assume Stability

Across saccades, small displacements of a visual target are harder to detect and their directions more difficult to discriminate than during steady fixation. Prominent theories of this effect, known as saccadic suppression of displacement, propose that it is due to a bias to assume object stability across saccades. Recent studies comparing the saccadic effect to masking effects suggest that suppression of displacement is not saccade-specific. Further evidence for this account is presented from two experiments where participants judged the size of displacements on a continuous scale in saccade and mask conditions, with and without blanking. Saccades and masks both reduced the proportion of correctly perceived displacements and increased the proportion of missed displacements. Blanking improved performance in both conditions by reducing the proportion of missed displacements. Thus, if suppression of displacement reflects a bias for stability, it is not a saccade-specific bias, but a more general stability assumption revealed under conditions of impoverished vision. Specifically, I discuss the potentially decisive role of motion or other transient signals for displacement perception. Without transients or motion, the quality of relative position signals is poor, and saccadic and mask-induced suppression of displacement reflects performance when the decision has to be made on these signals alone. Blanking may improve those position signals by providing a transient onset or a longer time to encode the pre-saccadic target position.

The eardrums move when the eyes move: A multisensory effect on the mechanics of hearing

Interactions between sensory pathways such as the visual and auditory systems are known to occur in the brain, but where they first occur is uncertain. Here, we show a multimodal interaction evident at the eardrum. Ear canal microphone measurements in humans (n = 19 ears in 16 subjects) and monkeys (n = 5 ears in three subjects) performing a saccadic eye movement task to visual targets indicated that the eardrum moves in conjunction with the eye movement. The eardrum motion was oscillatory and began as early as 10 ms before saccade onset in humans or with saccade onset in monkeys. These eardrum movements, which we dub eye movement-related eardrum oscillations (EMREOs), occurred in the absence of a sound stimulus. The amplitude and phase of the EMREOs depended on the direction and horizontal amplitude of the saccade. They lasted throughout the saccade and well into subsequent periods of steady fixation. We discuss the possibility that the mechanisms underlying EMREOs create eye movement-related binaural cues that may aid the brain in evaluating the relationship between visual and auditory stimulus locations as the eyes move.

Effect of Grouping, Segmentation, and Vestibular Stimulation on the Autokinetic Effect

We report some new observations on what could be regarded as the world’s simplest visual illusion—the autokinetic effect. When a single dim spot of light is viewed in a completely dark room, it moves vividly in random directions. During steady fixation, perhaps subtle eye movements cause the image to move and a failure to correct for this using eye movement command signals leads to motion perception. This is especially true because eye muscle fatigue can lead to miscalibration. However, if two dots are shown, they often move independently in different directions, which negate the eye movement theory. In addition, two lines defining a single cross sometimes uncouple and slide past each other and the fragments composing a hidden object move independently until they click in place and the whole object is perceived—implying that the illusion occurs relatively late in visual processing. Finally, the effect is modulated by vestibular stimulation; anchoring your sense of self may be a prerequisite for binding features into coherent objects.

The eardrums move when the eyes move: A multisensory effect on the mechanics of hearing

ABSTRACTInteractions between sensory pathways such as the visual and auditory systems are known to occur in the brain, but where they first occur is uncertain. Here we show a novel multimodal interaction evident at the eardrum. Ear canal microphone measurements in humans (n=19 ears in 16 subjects) and monkeys (n=5 ears in 3 subjects) performing a saccadic eye movement task to visual targets indicated that the eardrum moves in conjunction with the eye movement. The eardrum motion was oscillatory and began as early as 10 ms before saccade onset in humans or with saccade onset in monkeys. These eardrum movements, which we dub Eye Movement Related Eardrum Oscillations (EMREOs), occurred in the absence of a sound stimulus. The EMREOs’ amplitude and phase depended on the direction and horizontal amplitude of the saccade. They lasted throughout the saccade and well into subsequent periods of steady fixation. We discuss the possibility that the mechanisms underlying EMREOs create eye movement-related binaural cues that may aid the brain in evaluating the relationship between visual and auditory stimulus locations as the eyes move.SIGNIFICANCE STATEMENTThe peripheral hearing system contains several motor mechanisms that allow the brain to modify the auditory transduction process. Movements or tensioning of either the middle-ear muscles or the outer hair cells modify eardrum motion, producing sounds that can be detected by a microphone placed in the ear canal (e.g. as otoacoustic emissions). Here, we report a novel form of eardrum motion produced by the brain via these systems -- oscillations synchronized with and covarying with the direction and amplitude of saccades. These observations suggest that a vision-related process modulates the first stage of hearing. In particular, these eye-movement related eardrum oscillations may help the brain connect sights and sounds despite changes in the spatial relationship between the eyes and the ears.

Response Properties of Fixation Neurons and Their Location in the Frontal Eye Field in the Monkey

Electrical stimulation of the frontal eye field (FEF) has recently been reported to suppress the generation of saccades, which supports the idea that the FEF plays a role in maintaining attentive fixation. This study analyzed the activity of fixation neurons that discharged during fixation in the FEF in relation to visual fixation and saccades in trained monkeys. The neural activity of fixation neurons increased at the start of fixation and was maintained during fixation. When a fixation spot of light disappeared during steady fixation, different fixation neurons exhibited different categories of response, ranging from a decrease in activity to an increase in activity, indicating that there is a continuum of fixation neurons, from neurons with foveal visual-related activity to neurons with activity that is related to the motor act of fixating. Fixation neurons usually showed a decrease in their firing rate before the onset of visually guided saccades (Vsacs) and memory-guided saccades in any direction. The reduction in activity of fixation neurons nearly coincided with, or occurred slightly before, the increase in the activity of saccade-related movement neurons in the FEF in the same monkey. Although fixation neurons were scattered in the FEF, about two thirds of fixation neurons were concentrated in a localized area in the FEF at which electrical stimulation induced strong suppression of the initiation of Vsacs bilaterally. These results suggest that fixation neurons in the FEF are part of a suppression mechanism that could control the maintenance of fixation and the initiation of saccades.