Grey-out elimination: The roles of spatial waveform,frequency and phase

The retinal events of a saccadic eye movement were simulated by presenting to the stationary eye a blank field of variable duration bracketed in time by vertical gratings of the same average space luminance. The blank represents the saccadic “grey-out” and the gratings represent the clear retinal images of each fixational pause.The blank field fails to be be perceived when it is of short duration. This duration is dependent on the waveform and spatial frequency of the gratings and may be as long as 350 msec.The effect is termed “grey-out elimination” and could explain our failure to perceive the retinal smear produced by saccades.     

Changes in orientation discrimination at the time of saccadic eye movements

Perceptual performance has been known to change around the time of saccadic eye movement. In the current study, we measured the accuracy and sensitivity of orientation discrimination of bar stimuli presented during fixation and before saccadic eye movements. Human participants compared the orientations of the test and reference bar stimuli with the head erect in a two-interval forced choice task. For the targets presented during steady fixation, the accuracy and sensitivity of orientation discrimination were better near the cardinal than oblique axes, a perceptual anisotropy known as the oblique effect. For the targets presented during the 100 ms interval immediately before a saccade was executed, the anisotropy decreased mainly due to reduction in sensitivity for cardinal orientations. Directing attention to the goal location of the impending saccade emulated the saccadic effects on orientation discrimination for the targets at saccadic goal, suggesting that the saccadic effects on orientation discrimination are partly mediated by the shift of spatial attention that accompanies the saccade. These results were in line with the anti-oblique effect that perceptual judgment of motion direction along the oblique angle becomes relatively accurate for motion targets presented before saccadic eye movements [Lee, J., & Lee, C. (2005). Changes in visual motion perception before saccadic eye movements. Vision Research, 45(11), 1447-1457].     

Contrast sensitivity during the initiation of smooth pursuit eye movements

Eye movements challenge the perception of a stable world by inducing retinal image displacement. During saccadic eye movements visual stability is accompanied by a remapping of visual receptive fields, a compression of visual space and perceptual suppression. Here we explore whether a similar suppression changes the perception of briefly presented low contrast targets during the initiation of smooth pursuit eye movements. In a 2AFC design we investigated the contrast sensitivity for threshold-level stimuli during the initiation of smooth pursuit and during saccades. Pursuit was elicited by horizontal step-ramp and ramp stimuli. At any time from 200 ms before to 500 ms after pursuit stimulus onset, a blurred 0.3 deg wide horizontal line with low contrast just above detection threshold appeared for 10 ms either 2 deg above or below the pursuit trajectory. Observers had to pursue the moving stimulus and to indicate whether the target line appeared above or below the pursuit trajectory. In contrast to perceptual suppression effects during saccades, no pronounced suppression was found at pursuit onset for step-ramp motion. When pursuit was elicited by a ramp stimulus, pursuit initiation was accompanied by catch-up saccades, which caused saccadic suppression. Additionally, contrast sensitivity was attenuated at the time of pursuit or saccade stimulus onset. This attenuation might be due to an attentional deficit, because the stimulus required the focus of attention during the programming of the following eye movement.     

Motor signals in visual localization

We demonstrate a strong sensory-motor coupling in visual localization in which experimental modification of the control of saccadic eye movements leads to an associated change in the perceived location of objects. Amplitudes of saccades to peripheral targets were altered by saccadic adaptation, induced by an artificial step of the saccade target during the eye movement, which leads the oculomotor system to recalibrate saccade parameters. Increasing saccade amplitudes induced concurrent shifts in perceived location of visual objects. The magnitude of perceptual shift depended on the size and persistence of errors between intended and actual saccade amplitudes. This tight agreement between the change of eye movement control and the change of localization shows that perceptual space is shaped by motor knowledge rather than simply constructed from visual input.     

Selective attention and the active remapping of object features in trans-saccadic perception

When the same object is attended both before and after a saccadic eye movement, its visual features may be remapped to the new retinal position of the object. To further investigate the role of selective attention in trans-saccadic perception, the magnitude of the cross-saccadic tilt aftereffect was measured for both attended and unattended objects. The results show that both selective attention and saccadic eye movements influenced the magnitude of the tilt aftereffect, but in different ways. Dividing attention among multiple objects lead to a general decrease in the tilt aftereffect, independent of whether or not a saccade occurred. Making a saccade also resulted in a consistent reduction of the aftereffect, but this was due to incomplete transfer of form adaptation to the new retinal position. The influences of selective attention and saccadic remapping on the tilt aftereffect were independent and additive. These findings suggest that trans-saccadic perception is not limited to a single object but instead depends on the allocation of selective attention. Overall, the results are consistent with the hypothesis that the role of attention is to select salient objects, with trans-saccadic perception mechanisms acting to maintain information about those salient objects across eye movements.     

How presaccadic gratings modify postsaccadic modulation transfer function

Threshold modulations for sinusoidal gratings tachistoscopically presented on the fovea after a saccade were determined as a function of the spatial frequency of a peripheral grating at the saccade goal and compared with thresholds of the “resting eye” condition. The results show a clear enhancement for medium spatial frequency gratings (3.2 c/deg) with peripheral gratings of the same spatial frequency. Therefore an information transfer from periphery to centre induced by saccadic eye movements is suggested. Suppression of low spatial frequency gratings (0.5 c/deg) is found to be independent from additional peripheral gratings and interpreted as an effect of a central inhibitory process elicited by the saccadic motor command. Some aspects concerning saccadic suppression, refixation, and visual stability are considered.     

Non-veridical visual motion perception immediately after saccades.

It is widely assumed that combining the eye movement vector with the motion vector of the retinal image is both sufficient and necessary for recovering the direction and speed of visual motion. Here, we report that execution of a saccadic (rapid) eye movement in the dark systematically biased subsequent perceptual judgment of the direction of visual motion in the direction opposite to the saccade. This non-veridical motion perception reached a maximum immediately after saccade offset and then decayed in approximately equal to 100 ms. These results suggest that the oculomotor signal interacts with central mechanisms related to motion and possibly form perception, as well as spatial vision, as documented with mislocalization of visual objects at the time of saccades.     

Eye movement signals influence perception: evidence from the adaptation of reactive and volitional saccades

Information about upcoming saccadic eye movements is used to orient visuo-spatial attention across the visual field. Different eye movement signals (intended or actual) could be used according to the intentionality of the saccade in preparation (Reactive or Volitional), and can be dissociated by saccadic adaptation. Gap 0 and overlap paradigms were contrasted to elicit the two saccade populations with different latencies and an asymmetric transfer of saccadic adaptation. Preparation of both saccade types caused a concomitant shift in the attentional focus (indexed by relative perceptual performance) to the actual, not intended, eye position. The attentional shift emerged progressively, earlier for V-saccades but reaching a maximal level around saccade onset for both saccade types. These results suggest that information about actual eye movements mediates the pre-saccadic shift of attention.     

Movement perception during voluntary saccadic eye movements

In order to elucidate the stabilization mechanism of the visual perception during voluntary eye movements the perception of a moving object during a eye saccade was investigated in human subjects. The results were analyzed in terms of velocity and displacement perception channels. The experimental results indicate that during voluntary saccades there is no suppression of movement perception, and, on the other hand, that in darkness the direction of the perceived movement is that of the saccade whereas in light the direction of the objective movement prevails. The latter discrepancy can be explained by assuming that the perceptual analysis of movement during the saccade occurs in the displacement channel and that this channel is provided with a channel evaluation mechanism controlled by the oculomotor centers.     

Modification of eye movements by instantaneous changes in the velocity of visual targets

The question of how information concerning the motion of targets becomes manifested in eye movements was answered in the following way. Monkeys, previously taught to track visual targets, were exposed to targets whose velocity was instantaneously changed at varying intervals after the onset of target movement. Both saccadic and smooth pursuit eye movements could be altered by changes in target velocity introduced during the first 90 msec of target movement. A change in the velocity of the target could affect the size of a saccade occurring only 50 msec later. Saccadic eye movements occurred with intersaccadic intervals as short as 20 msec. These results indicate that both saccadic and smooth pursuit eye movements are based on continuously processed information.     

Conjugate ocular oscillations during shifts of the direction and depth of visual fixation.

PURPOSE: To characterize dynamic properties of combined saccade-vergence eye movements that occur as the point of visual fixation is shifted between objects lying in different directions and at different depths. METHODS: Using the scleral search-coil technique, eye movements were measured in 10 normal subjects as they made voluntary, disjunctive gaze shifts comprising a range of saccades and vergence movements. RESULTS: By analyzing eye acceleration records, the authors identified small-amplitude (0.2-0.7 degrees), high-frequency (23-33 Hz), conjugate horizontal oscillations of the eyes during the vergence movement that followed the initial saccade. When the shift of the fixation point required a large vergence component (17 degrees , every subject showed these oscillations; they were present in approximately a third of responses. Approximately 5% of responses showed oscillations that had horizontal and vertical components. Oscillations were less prominent with shifts that had smaller vergence components and were absent after saccades made between targets located at optical infinity. CONCLUSIONS: These findings suggest that a common mechanism gates both the saccadic and vergence components of disjunctive gaze shifts, a likely candidate being the pontine omnipause neurons. When a saccade is immediately followed by a prolonged vergence movement, the omnipause neurons remain silent, leading to small-amplitude saccadic oscillations. Shifts in the point of visual fixation that require a large vergence movement may be a useful experimental strategy to induce saccadic oscillations.     

Neuronal mechanisms of visual stability

Human vision is stable and continuous in spite of the incessant interruptions produced by saccadic eye movements. These rapid eye movements serve vision by directing the high resolution fovea rapidly from one part of the visual scene to another. They should detract from vision because they generate two major problems: displacement of the retinal image with each saccade and blurring of the image during the saccade. This review considers the substantial advances in understanding the neuronal mechanisms underlying this visual stability derived primarily from neuronal recording and inactivation studies in the monkey, an excellent model for systems in the human brain. For the first problem, saccadic displacement, two neuronal candidates are salient. First are the neurons in frontal and parietal cortex with shifting receptive fields that provide anticipatory activity with each saccade and are driven by a corollary discharge. These could provide the mechanism for a retinotopic hypothesis of visual stability and possibly for a transsaccadic memory hypothesis, The second neuronal mechanism is provided by neurons whose visual response is modulated by eye position (gain field neurons) or are largely independent of eye position (real position neurons), and these neurons could provide the basis for a spatiotopic hypothesis. For the second problem, saccadic suppression, visual masking and corollary discharge are well established mechanisms, and possible neuronal correlates have been identified for each.     

Binocular eye movements during accommodative vergence

Binocular eye position was monitored by the photoelectric technique during accommodative vergence. Contrary to previous reports indicating that accommodative vergence was a uniocular phenomenon, without exception, binocular accommodative vergence movements were recorded. The total vergence amplitude in the viewing eye was reduced, on the average, by approximately 88% with respect to the vergence movement measured in the covered eye. Some saccadic eye movements that occurred during vergence movements were likewise reduced in amplitude in the viewing eye by up to 20%. Smooth eye movements were utilized to counteract the vergence movement in the viewing eye. This smooth movement alone, or in conjunction with a late saccade, returned the eye to the target and helped to maintain the retinal image of the target coincident with the foveal center for the duration of the accommodative vergence movement. Thus, there appears to be a fixation-holding mechanism which produced a general attenuation of both vergence and some saccadic movements in the viewing eye. Although this control strategy produced violations of Hering's law with respect to the magnitude of the movements in the eyes but not with respect to the direction of the movement, it was implemented in the interest of retaining the target within the sensitive foveal region.     

Spatio-temporal topography of saccadic overestimation of time

Rapid eye movements (saccades) induce visual misperceptions. A number of studies in recent years have investigated the spatio-temporal profiles of effects like saccadic suppression or perisaccadic mislocalization and revealed substantial functional similarities. Saccade induced chronostasis describes the subjective overestimation of stimulus duration when the stimulus onset falls within a saccade. In this study we aimed to functionally characterize saccade induced chronostasis in greater detail. Specifically we tested if chronostasis is influenced by or functionally related to saccadic suppression. In a first set of experiments, we measured the perceived duration of visual stimuli presented at different spatial positions as a function of presentation time relative to the saccade. We further compared perceived duration during saccades for isoluminant and luminant stimuli. Finally, we investigated whether or not saccade induced chronostasis is dependent on the execution of a saccade itself. We show that chronostasis occurs across the visual field with a clear spatio-temporal tuning. Furthermore, we report chronostasis during simulated saccades, indicating that spurious retinal motion induced by the saccade is a prime origin of the phenomenon.     

Age effects on saccadic suppression of luminance and color

Saccadic eye movements modulate visual perception: they initiate and terminate high acuity vision at a certain location in space, but before and during their execution visual contrast sensitivity is strongly attenuated for 100 to 200 ms. Transient perisaccadic perceptual distortions are assumed to be an important mechanism to maintain visual stability. Little is known about age effects on saccadic suppression, even though for healthy adults other major age-related changes are well documented, like a decrease of visual contrast sensitivity for intermediate and high spatial frequencies or an increase of saccade latencies. Here, we tested saccadic suppression of luminance and isoluminant chromatic flashes in 100 participants from eight to 78 years. To estimate the effect of saccadic suppression on contrast sensitivity, we used a two-alternative forced choice (2AFC) design and an adaptive staircase procedure to modulate the luminance or chromatic contrast of a flashed detection target during fixation and 15 ms after saccade onset. The target was a single horizontal luminance or chromatic line flashed 2° above or below the fixation or saccade target. Compared to fixation, average perisaccadic contrast sensitivity decreased significantly by 66% for luminance and by 36% for color. A significant correlation was found for the strength of saccadic suppression of luminance and color. However, a small age effect was found only for the strength of saccadic suppression of luminance, which increased from 64% to 70% from young to old age. We conclude that saccadic suppression for luminance and color is present in most participants independent of their age and that mechanisms of suppression stay relatively stable during healthy aging.     

Landmarks facilitate visual space constancy across saccades and during fixation

It has been demonstrated that visual objects that are present after saccadic eye movements act as landmarks for the localization of stimuli across saccades, facilitating space constancy (Deubel, 2004). We here study the temporal conditions under which landmark effects occur after saccadic eye movements, and during fixation. Two small objects were presented 6° in the periphery, one above the other. Observers saccaded to the space between them. One of the objects disappeared during the saccade and reappeared with a variable delay during or after the saccade. At the same time either that object or the continuously present one jumped by 1°. The observer’s task was to decide which object had moved. The results revealed a strong bias to assign movement to the object that was blanked, regardless of which actually moved. If both objects were blanked, the one that was blanked for a shorter time tended to be seen as stable. The effects were stronger as the onset asynchrony between the stimuli increased. Surprisingly, analogous though weaker effects occurred during visual fixation, suggesting that similar visual mechanisms relying on visual landmarks operate both across saccades and during fixation.     

Horizontal saccade disconjugacy in strabismic monkeys

PURPOSE: Previous studies have shown that binocular coordination during saccadic eye movement is affected in humans with large strabismus. The purpose of this study was to examine the conjugacy of saccadic eye movements in monkeys with sensory strabismus. METHODS: The authors recorded binocular eye movements in four strabismic monkeys and one unaffected monkey. Strabismus was induced by first occluding one eye for 24 hours, switching the occluder to the fellow eye for the next 24 hours, and repeating this pattern of daily alternating monocular occlusion for the first 4 to 6 months of life. Horizontal saccades were measured during monocular viewing when the animals were 2 to 3 years of age. RESULTS: Horizontal saccade testing during monocular viewing showed that the amplitude of saccades in the nonviewing eye was usually different from that in the viewing eye (saccade disconjugacy). The amount of saccade disconjugacy varied among animals as a function of the degree of ocular misalignment as measured in primary gaze. Saccade disconjugacy also increased with eccentric orbital positions of the nonviewing eye. If the saccade disconjugacy was large, there was an immediate postsaccadic drift for less than 200 ms. The control animal showed none of these effects. CONCLUSIONS: As do humans with large strabismus, strabismic monkey display disconjugate saccadic eye movements. Saccade disconjugacy varies with orbital position and increases as a function of ocular misalignment as measured in primary gaze. This type of sensory-induced strabismus serves as a useful animal model to investigate the neural or mechanical factors responsible for saccade disconjugacy observed in humans with strabismus.     

Attenuation of perceived motion smear during vergence and pursuit tracking

When the eyes move, the images of stationary objects sweep across the retina. Despite this motion of the retinal image and the substantial integration of visual signals across time, physically stationary objects typically do not appear to be smeared during eye movements. Previous studies indicated that the extent of perceived motion smear is smaller when a stationary target is presented during pursuit or saccadic eye movements than when comparable motion of the retinal image occurs during steady fixation. In this study, we compared the extent of perceived motion smear for a stationary target during smooth pursuit and vergence eye movements with that for a physically moving target during fixation. For a target duration of 100 ms or longer, perceived motion smear is substantially less when the motion of the retinal image results from vergence or pursuit eye movements than when it results from the motion of a target during fixation. The reduced extent of perceived motion smear during eye movements compared to fixation cannot be accounted for by different spatio-temporal interactions between visual targets or by unequal attention to the moving test spot under these two types of conditions. We attribute the highly similar attenuation of perceived smear during vergence and pursuit to a comparable action of the extra-retinal signals for disjunctive and conjugate eye movements.     

Saccadic suppression of low-level motion

We measured the detection of motion before, during and after a saccade to explore the effects of a saccade on motion perception. To isolate the low-level motion mechanism, the stimulus was a random-dot field displaced by small distance (0.3 deg) within a stationary frame. The displacement signaled motion clearly if eyes were fixated, but for the displacement during a saccade, motion was not detected whether the displacement was defined in spatial coordinates (expt 1) or in retinal coordinates (expt 2). Since motion could be seen with ISIs longer than the duration of a saccade (expt 3), the suppression cannot be attributed to visual loss during the saccade. Experiment 3 also showed that motion was never seen for a displacement that occurred during a saccade, even though the random dots were replaced by a uniform field during the eye movement thereby eliminating any masking effect of the sweep of the image across the retina. The purpose of the saccadic suppression of motion may be to block out unreliable motion signals that would be produced by a saccade. Since saccade distances are very often greater than the maximum distance over which the low-level motion mechanism can produce accurate direction discrimination for fine textures, motion signals would generally indicate false directions if they were not suppressed.     

Spatial attention and latencies of saccadic eye movements.

Recent theories of visual attention, such as the oculomotor readiness theory of Klein (1980) (Does oculomotor readiness mediate cognitive control of the visual attention. In: R. Nickerson, Attention and performance, Hillsdale: Erlbaum), the premotor theory of Rizzolati (1983) (Mechanisms of selective attention in mammals. In: J.P. Ewart, R.R. Capranica, D.J. Ingle, Advances in vertebrate Neuroethology (pp. 261-297). New York: Plenum) and the sequential attention theory of Henderson (1992) (Visual attention and eye movement control during reading and scene perception. In K. Rayner, Eye movements and visual cognition (260-283). New York: Springer-Verlag), propose a link between shifts in spatial attention and the generation of saccadic eye movements. In this paper we show that a winner-take-all model of spatial attention, combined with a simple model for the link between attention and eye movements, can account for the variation in saccadic latency observed in many oculomotor phenomena. These phenomena include the gap effect (Saslow M.G. (1967). Effects of components of displacement-step stimuli upon latency for saccadic eye movement. Journal of the Optical Society of America, 57, 1024-1029), the effect of target jumps on saccadic latency (Becker W. & Jurgens R. (1979). An analysis of the saccadic system by means of double step stimuli. Vision Research, 19, 967-983), the increase of saccadic latency as target eccentricity drops (Kalesnykas R.P. & Hallett P.E. (1994). Retinal eccentricity and the latency of eye saccades. Vision Research, 34, 517-531), and the modulation of saccadic accuracy using target predictability and saccadic latency (Co?ffé C. & O'Regan J.K. (1987). Reducing the influence of non-target stimuli on saccade accuracy: predictability and latency effects. Vision Research, 27 (2), 227-240).