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2007 |
Samuel B. Hutton; Brendan S. Weekes Low frequency rTMS over posterior parietal cortex impairs smooth pursuit eye tracking Journal Article In: Experimental Brain Research, vol. 183, no. 2, pp. 195–200, 2007. @article{Hutton2007, The role of the posterior parietal cortex in smooth pursuit eye movements remains unclear. We used low frequency repetitive transcranial magnetic stimulation (rTMS) to study the cognitive and neural systems involved in the control of smooth pursuit eye movements. Eighteen participants were tested on two separate occasions. On each occasion we measured smooth pursuit eye tracking before and after 6 min of 1 Hz rTMS delivered at 90% of motor threshold. Low frequency rTMS over the posterior parietal cortex led to a significant reduction in smooth pursuit velocity gain, whereas rTMS over the motor cortex had no effect on gain. We conclude that low frequency offline rTMS is a potentially useful tool with which to explore the cortical systems involved in oculomotor control. |
Stephanie Jainta; Jörg Hoormann; W. Jaschinski Objective and subjective measures of vergence step responses Journal Article In: Vision Research, vol. 47, no. 26, pp. 3238–3246, 2007. @article{Jainta2007, Dichoptic nonius lines are used for subjectively (psychophysically) measuring vergence states, but they have been questioned as valid indicators of vergence eye position. In a mirror-stereoscope, we presented convergent and divergent step-stimuli and estimated the vergence response with nonius lines flashed at fixed delays after the disparity step stimulus. For each delay, an adaptive psychophysical procedure was run to determine the physical nonius offset required for subjective alignment; these vergence states were compared with objective eye movement recordings. Between both measures of initial vergence, we calculated the maximal cross-correlation coefficient: the median in our sample was about 0.9 for convergence and divergence, suggesting a good agreement. Relative to the objective measures, the subjective method revealed a smaller vergence velocity and a larger vergence response in the final phase of the response, but both measures were well correlated. The dynamic nonius test is therefore considered to be useful to relatively evaluate a subject's ability in disparity vergence. |
Wolfgang Jaschinski; Stephanie Jainta; Jörg Hoormann; Nina Walper Objective vs subjective measurements of dark vergence Journal Article In: Ophthalmic and Physiological Optics, vol. 27, no. 1, pp. 85–92, 2007. @article{Jaschinski2007, Dark vergence is a resting position of vergence (tonic vergence), measured in a dark visual field to eliminate fusional, accommodative, and proximal stimuli. The vergence resting position is relevant for measures of phoria and fixation disparity. Dark vergence differs reliably among subjects: the average subject converges at a viewing distance of about 1 m, while the inter-individual range is from infinity to about 40 cm. In previous research, dark vergence was measured subjectively, i.e. observers adjusted the horizontal offset of dichoptically presented nonius targets to perceived alignment. Results of such subjective vergence tests do not necessarily agree with those of the objective measurements of eye position with eye trackers. Therefore, we made simultaneous subjective and objective measurements of dark vergence and found similar results with both methods in repeated tests in two sessions. Thus, the nonius test is sufficient for a subjective estimation of dark vergence. |
Andre Kaminiarz; Bart Krekelberg; Frank Bremmer Localization of visual targets during optokinetic eye movements Journal Article In: Vision Research, vol. 47, no. 6, pp. 869–878, 2007. @article{Kaminiarz2007, We investigated localization of brief visual targets during reflexive eye movements (optokinetic nystagmus). Subjects mislocalized these targets in the direction of the slow eye movement. This error decreased shortly before a saccade and temporarily increased afterwards. The pattern of mislocalization differs markedly from mislocalization during voluntary eye movements in the presence of visual references, but (spatially) resembles mislocalization during voluntary eye movements in darkness. Because neither reflexive eye movements nor voluntary eye movements in darkness have explicit (visual) goals, these data support the view that visual goals support perceptual stability as an important link between pre- and post-saccadic scenes. |
Stephen J. Kerrigan; John F. Soechting Anisotropies in the gain of smooth pursuit during two-dimensional tracking as probed by brief perturbations Journal Article In: Experimental Brain Research, vol. 180, no. 3, pp. 435–448, 2007. @article{Kerrigan2007, Previous investigations suggest the gain of smooth pursuit is directionally anisotropic and is regulated in a task-dependent manner. Smooth pursuit is also known to be influenced by expectations concerning the target's motion, but the role of such expectations in modulating feedback gain is not known. In the present work, the gain of smooth pursuit was probed by applying brief perturbations to quasi-predictable two-dimensional target motion at multiple time points. The target initially moved in a straight line, then followed the circumference of a circle for distances ranging between 180 degrees and 270 degrees . Finally, the path reverted to linear motion. Perturbations consisted of a pulse of velocity 50 or 100 ms in duration, applied in one of eight possible directions. They were applied at the onset of the curve or after the target had traversed an arc of 45 degrees or 90 degrees . Pursuit gain was measured by computing the average amplitude of the response in smooth pursuit velocity over a 100 ms interval. To do so we used a coordinate system defined by the motion of the target at the onset of the perturbation, with directions tangential and normal to the path. Responses to the perturbations had two components: one that was modulated with the direction of the perturbation and one that was directionally nonspecific. For the directional response, on average the gain in the normal direction was slightly larger than the gain in the tangential direction, with a ratio ranging from 1.0 to 1.3. The directionally nonspecific response, which was more prominent for perturbations at curve onset or at 90 degrees , consisted of a transient decrease in pursuit speed. Perturbations applied at curve onset also delayed the tracking of the curved target motion. |
K. Konigs; J. Knoll; Frank Bremmer Localisation of auditory targets during optokinetic nystagmus Journal Article In: Perception, vol. 36, pp. 1507–1513, 2007. @article{Konigs2007, Previous studies have shown that the perceived location of visual stimuli briefly flashed during smooth pursuit, saccades, or optokinetic nystagmus (OKN) is not veridical. We investigated whether these mislocalisations can also be observed for brief auditory stimuli presented during OKN. Experiments were carried out in a lightproof sound-attenuated chamber. Participants performed eye movements elicited by visual stimuli. An auditory target (white noise) was presented for 5 ms. Our data clearly indicate that auditory targets are mislocalised during reflexive eye movements. OKN induces a shift of perceived location in the direction of the slow eye movement and is modulated in the temporal vicinity of the fast phase. The mislocalisation is stronger for look- as compared to stare-nystagmus. The size and temporal pattern of the observed mislocalisation are different from that found for visual targets. This suggests that different neural mechanisms are at play to integrate oculomotor signals and information on the spatial location of visual as well as auditory stimuli. |
2006 |
Dirk Kerzel; M. Pilar Aivar; Nathalie E. Ziegler; Eli Brenner Mislocalization of flashes during smooth pursuit hardly depends on the lighting conditions Journal Article In: Vision Research, vol. 46, no. 6-7, pp. 1145–1154, 2006. @article{Kerzel2006, Targets that are briefly flashed during smooth pursuit eye movements are mislocalized in the direction of motion (forward shift) and away from the fovea (spatial expansion). Hansen [Hansen, R. M. (1979). Spatial localization during pursuit eye movements. Vision Research 19(11), 1213-1221] reported that these errors are not present for fast motor responses in the dark, whereas Rotman et al. [Rotman, G., Brenner, E., Smeets, J. B. (2004). Quickly tapping targets that are flashed during smooth pursuit reveals perceptual mislocalizations. Experimental Brain Research 156(4), 409-414] reported that they are present for fast motor responses in the light. To evaluate whether the lighting conditions are the critical factor, we asked observers to point to the positions of flashed objects during smooth pursuit either in the dark or with the room lights on. In a first experiment, the flash, which could appear at 1 of 15 different positions, was always shown when the eye had reached a certain spatial position. We found a forward bias and spatial expansion that were independent of the target and ambient luminance. In a second experiment, the flash was always shown at the same retinal position, but the spatial position of the eye at the moment of flash presentation was varied. In this case we found differences between the luminance conditions, in terms of how the errors depended on the velocity and position on the trajectory. We also found specific conditions in which people did not mislocalize the target in the direction of pursuit at all. These findings may account for the above-mentioned discrepancy. We conclude that although the lighting conditions do influence the localization errors under some circumstances, it is certainly not so that such errors are absent whenever the experiment is conducted in the dark. |
Mark F. Lenzenweger; Gillian A. O'Driscoll Smooth pursuit eye movement and schizotypy in the community Journal Article In: Journal of Abnormal Psychology, vol. 115, no. 4, pp. 779–786, 2006. @article{Lenzenweger2006, Deficits in smooth pursuit eye movements are well documented in schizophrenia and schizotypic psychopathology. The status of eye tracking dysfunction (ETD) as an endophenotype for schizophrenia liability is relatively robust. However, the relation of ETD to schizophrenia-related deviance in the general population has not been confirmed. This study examined smooth pursuit eye tracking and schizotypal personality features in the general population. Smooth pursuit eye movement and schizotypal features were measured in 300 adult community subjects. The sample included both sexes, subjects with a wide age and educational range, and subjects with no prior history of psychosis. Primary outcome measures were peak gain (eye velocity/target velocity), catch-up saccade rate, and schizotypal feature scores. Total schizotypal features were significantly associated with decreased peak gain and were associated at the trend level with increased catch-up saccade rate. These associations were essentially unchanged after controlling for age, sex, and intellectual level effects. These data confirm a hypothesized association between schizotypal features and poorer eye tracking performance (principally, peak gain) in the general population as well as support the conceptualization of ETD as an endophenotype for schizophrenia liability. |
Jan L. Souman; Ignace T. C. Hooge; Alexander H. Wertheim Frame of reference transformations in motion perception during smooth pursuit eye movements Journal Article In: Journal of Computational Neuroscience, vol. 20, no. 1, pp. 61–76, 2006. @article{Souman2006, Smooth pursuit eye movements change the retinal image velocity of objects in the visual field. In order to change from a retinocentric frame of reference into a head-centric one, the visual system has to take the eye movements into account. Studies on motion perception during smooth pursuit eye movements have measured either perceived speed or perceived direction during smooth pursuit to investigate this frame of reference transformation, but never both at the same time. We devised a new velocity matching task, in which participants matched both perceived speed and direction during fixation to that during pursuit. In Experiment 1, the velocity matches were determined for a range of stimulus directions, with the head-centric stimulus speed kept constant. In Experiment 2, the retinal stimulus speed was kept approximately constant, with the same range of stimulus directions. In both experiments, the velocity matches for all directions were shifted against the pursuit direction, suggesting an incomplete transformation of the frame of reference. The degree of compensation was approximately constant across stimulus direction. We fitted the classical linear model, the model of Turano and Massof (2001) and that of Freeman (2001) to the velocity matches. The model of Turano and Massof fitted the velocity matches best, but the differences between de model fits were quite small. Evaluation of the models and comparison to a few alternatives suggests that further specification of the potential effect of retinal image characteristics on the eye movement signal is needed. |
Jan L. Souman; Ignace T. C. Hooge; Alexander H. Wertheim Localization and motion perception during smooth pursuit eye movements Journal Article In: Experimental Brain Research, vol. 171, no. 4, pp. 448–458, 2006. @article{Souman2006a, We investigated the relationship between compensation for the effects of smooth pursuit eye movements in localization and motion perception. Participants had to indicate the perceived motion direction, the starting point and the end point of a vertically moving stimulus dot presented during horizontal smooth pursuit. The presentation duration of the stimulus was varied. From the indicated starting and end points, the motion direction was predicted and compared with the actual indicated directions. Both the directions predicted from localization and the indicated directions deviated from the physical directions, but the errors in the predicted directions were larger than those in the indicated directions. The results of a control experiment, in which the same tasks were performed during fixation, suggest that this difference reflects different transformations from a retinocentric to a head-centric frame of reference. This difference appears to be mainly due to an asymmetry in the effect of retinal image motion direction on localization during smooth pursuit. |
Miriam Spering; Karl R. Gegenfurtner; Dirk Kerzel Distractor interference during smooth pursuit eye movements Journal Article In: Journal of Experimental Psychology: Human Perception and Performance, vol. 32, no. 5, pp. 1136–1154, 2006. @article{Spering2006, When 2 targets for pursuit eye movements move in different directions, the eye velocity follows the vector average (S. G. Lisberger & V. P. Ferrera, 1997). The present study investigates the mechanisms of target selection when observers are instructed to follow a predefined horizontal target and to ignore a moving distractor stimulus. Results show that at 140 ms after distractor onset, horizontal eye velocity is decreased by about 25%. Vertical eye velocity increases or decreases by 1 degrees /s in the direction opposite from the distractor. This deviation varies in size with distractor direction, velocity, and contrast. The effect was present during the initiation and steady-state tracking phase of pursuit but only when the observer had prior information about target motion. Neither vector averaging nor winner-take-all models could predict the response to a moving to-be-ignored distractor during steady-state tracking of a predefined target. The contributions of perceptual mislocalization and spatial attention to the vertical deviation in pursuit are discussed. |
Minah Suh; Sambrita Basu; Rachel Kolster; Ranjeeta Sarkar; Bruce D. McCandliss; Jamshid Ghajar Increased oculomotor deficits during target blanking as an indicator of mild traumatic brain injury Journal Article In: Neuroscience Letters, vol. 410, no. 3, pp. 203–207, 2006. @article{Suh2006, Given the susceptibility of cerebellar-cortical tracts to shearing injury from traumatic brain injury (TBI), we investigated impairment in the generation of predictive eye movements and its relationship to cognitive deficits in mild TBI patients using a smooth pursuit target-blanking paradigm. Compared to a target-tracking paradigm without blanking, this paradigm more greatly necessitates the generation of predictive eye movements, which are subserved by brain regions involved in cognitive processing. Mild TBI patients showed impaired prediction of target trajectories during target blanking, demonstrated by generation of saccades at earlier and more variable time points, as well as greater and more variable oculomotor error compared to controls. In addition, California Verbal Learning Test (CVLT-II) scores related to working memory, learning, and executive function were more highly correlated with oculomotor variability during target blanking than during target tracking. Our results suggest that a disruption of cerebellar-cortical connections in TBI may account for both oculomotor and cognitive impairment, and that measures of predictive eye movements during target blanking may be a sensitive metric of cognitive deficits after mild TBI. |
Minah Suh; Rachel Kolster; Ranjeeta Sarkar; Bruce D. McCandliss; Jamshid Ghajar Deficits in predictive smooth pursuit after mild traumatic brain injury Journal Article In: Neuroscience Letters, vol. 401, no. 1-2, pp. 108–113, 2006. @article{Suh2006a, Given that even mild traumatic brain injury (TBI) may produce extensive diffuse axonal injury (DAI), we hypothesized that mild TBI patients would show deficits in predictive smooth pursuit eye movements (SPEM), associated with impaired cognitive functions, as these processes are dependent on common white matter connectivity between multiple cerebral and cerebellar regions. The ability to predict target trajectories during SPEM was investigated in 21 mild TBI patients using a periodic sinusoidal paradigm. Compared to 26 control subjects, TBI patients demonstrated decreased target prediction. TBI patients also showed increased eye position error and variability of eye position, which correlated with decreased target prediction. In all subjects, average target prediction, eye position error and eye position variability correlated with scores related to attention and executive function on the California Verbal Learning Test (CVLT-II). However, there were no differences between TBI and control groups in average eye gain or intra-individual eye gain variability, or in performance on the Wechsler Abbreviated Scale of Intelligence (WASI), suggesting that the observed deficits did not result from general oculomotor impairment or reduced IQ. The correlation between SPEM performance and CVLT-II scores suggests that predictive SPEM may be a sensitive assay of cognitive functioning, including attention and executive function. This is the first report to our knowledge that TBI patients show impaired predictive SPEM and eye position variability, and that these impairments correlate with cognitive deficits. |
Benjamin W. Tatler; Roland J. Baddeley; Benjamin T. Vincent The long and the short of it: Spatial statistics at fixation vary with saccade amplitude and task Journal Article In: Vision Research, vol. 46, no. 12, pp. 1857–1862, 2006. @article{Tatler2006, We recorded over 90,000 saccades while observers viewed a diverse collection of natural images and measured low level visual features at fixation. The features that discriminated between where observers fixated and where they did not varied considerably with task, and the length of the preceding saccade. Short saccades (<8°) are image feature dependent, long are less so. For free viewing, short saccades target high frequency information, long saccades are scale-invariant. When searching for luminance targets, saccades of all lengths are scale-invariant. We argue that models of saccade behaviour must account not only for task but also for saccade length and that long and short saccades are targeted differently. |
Martin Rolfs; Jochen Laubrock; Reinhold Kliegl Shortening and prolongation of saccade latencies following microsaccades Journal Article In: Experimental Brain Research, vol. 169, no. 3, pp. 369–376, 2006. @article{Rolfs2006, When the eyes fixate at a point in a visual scene, small saccades rapidly shift the image on the retina. The effect of these microsaccades on the latency of subsequent large-scale saccades may be twofold. First, microsaccades are associated with an enhancement of visual perception. Their occurrence during saccade target perception could, thus, decrease saccade latencies. Second, microsaccades are likely to indicate activity in fixation-related oculomotor neurons. These represent competitors to saccade-related cells in the interplay of gaze holding and shifting. Consequently, an increase in saccade latencies would be expected after microsaccades. Here, we present evidence for both aspects of microsaccadic impact on saccade latency. In a delayed response task, participants made saccades to visible or memorized targets. First, microsaccade occurrence up to 50 ms before target disappearance correlated with 18 ms (or 8%) faster saccades to memorized targets. Second, if microsaccades occurred shortly (i.e., <150 ms) before a saccade was required, mean saccadic reaction time in visual and memory trials was increased by about 40 ms (or 16%). Hence, microsaccades can have opposite consequences for saccade latencies, pointing at a differential role of these fixational eye movements in the preparation of saccade motor programs. |
Ralf Engbert; Konstantin Mergenthaler Microsaccades are triggered by low retinal image slip Journal Article In: Proceedings of the National Academy of Sciences, vol. 103, no. 18, pp. 7192–7197, 2006. @article{Engbert2006, Even during visual fixation of a stationary target, our eyes perform rather erratic miniature movements, which represent a random walk. These "fixational" eye movements counteract perceptual fading, a consequence of fast adaptation of the retinal receptor systems to constant input. The most important contribution to fixational eye movements is produced by microsaccades; however, a specific function of microsaccades only recently has been found. Here we show that the occurrence of microsaccades is correlated with low retinal image slip approximately 200 ms before microsaccade onset. This result suggests that microsaccades are triggered dynamically, in contrast to the current view that microsaccades are randomly distributed in time characterized by their rate-of-occurrence of 1 to 2 per second. As a result of the dynamic triggering mechanism, individual microsaccade rate can be predicted by the fractal dimension of trajectories. Finally, we propose a minimal computational model for the dynamic triggering of microsaccades. |
Casper J. Erkelens Coordination of smooth pursuit and saccades Journal Article In: Vision Research, vol. 46, no. 1-2, pp. 163–170, 2006. @article{Erkelens2006, Smooth pursuit and saccades are two components of tracking eye movements. Their coordination has usually been studied by investigating latencies of pursuit onset in response to a moving target appearing simultaneously with the disappearance of the stationary fixation target. The general finding from such studies has been that latencies of saccades and pursuit are different and reflect independent processes. We discuss several limitations of the used targets. In this paper, we study latencies of saccades and smooth pursuit in response to a moving target that overlaps in time with a pursued moving target. We find that saccades and pursuit changes are synchronized. Furthermore, pursuit changes are made fast. Directional changes occur almost entirely within the accompanying saccade. To explain the results we hypothesize a two-stage mechanism for the coordinated generation of saccades and pursuit. |
Kai Essig; Marc Pomplun; Helge Ritter A neural network for 3D gaze recording with binocular eye trackers Journal Article In: International Journal of Parallel, Emergent and Distributed Systems, vol. 21, no. 2, pp. 79–95, 2006. @article{Essig2006, Using eye tracking for the investigation of visual attention has become increasingly popular during the last few decades. Nevertheless, only a small number of eye tracking studies have employed 3D displays, although such displays would closely resemble our natural visual environment. Besides higher cost and effort for the experimental setup, the main reason for the avoidance of 3D displays is the problem of computing a subject's current 3D gaze position based on the measured binocular gaze angles. The geometrical approaches to this problem that have been studied so far involved substantial error in the measurement of 3D gaze trajectories. In order to tackle this problem, we developed an anaglyph-based 3D calibration procedure and used a well-suited type of artificial neural network—a parametrized self-organizing map (PSOM)—to estimate the 3D gaze point from a subject's binocular eye-position data. We report an experiment in which the accuracy of the PSOM gaze-point estimation is compared to a geometrical solution. The results show that the neural network approach produces more accurate results than the geometrical method, especially for the depth axis and for distant stimuli. |
Danny Gagnon Transcranial magnetic stimulation of frontal oculomotor regions during smooth pursuit Journal Article In: Journal of Neuroscience, vol. 26, no. 2, pp. 458–466, 2006. @article{Gagnon2006, Both the frontal eye fields (FEFs) and supplementary eye fields (SEFs) are known to be involved in smooth pursuit eye movements. It has been shown recently that stimulation of the smooth-pursuit area of the FEF [frontal pursuit area (FPA)] in monkey increases the pursuit response to unexpected changes in target motion during pursuit. In the current study, we applied transcranial magnetic stimulation (TMS) to the FPA and SEF in humans during sinusoidal pursuit to assess its effects on the pursuit response to predictable, rather than unexpected, changes in target motion. For the FPA, we found that TMS applied immediately before the target reversed direction increased eye velocity in the new direction, whereas TMS applied in mid-cycle, immediately before the target began to slow, decreased eye velocity. For the SEF, TMS applied at target reversal increased eye velocity in the new direction but had no effect on eye velocity when applied at mid-cycle. TMS of the control region (leg region of the somatosensory cortex) did not affect eye velocity at either point. Previous stimulation studies of FPA during pursuit have suggested that this region is involved in controlling the gain of the transformation of visual signals into pursuit motor commands. The current results suggest that the gain of the transformation of predictive signals into motor commands is also controlled by the FPA. The effect of stimulation of the SEF is distinct from that of the FPA and suggests that its role in sinusoidal pursuit is primarily at the target direction reversal. |
P. U. Tse; G. P. Caplovitz; P. -J. Hsieh Microsaccade directions do not predict directionality of illusory brightness changes of overlapping transparent surfaces Journal Article In: Vision Research, vol. 46, no. 22, pp. 3823–3830, 2006. @article{Tse2006a, Tse (2005) recently introduced a new class of illusory brightness changes where shifts of attention lead to shifts in perceived brightness across overlapping, transparent figures, under conditions of visual fixation. In the absence of endogenous attentional shifts, illusory brightness changes appear to shift from figure to figure spontaneously, much as occurs in other multistable phenomena. The goal of the present research is to determine whether fixational microsaccades are correlated with perceived brightness changes. It has recently been demonstrated that microsaccades can reveal the direction of covert attentional shifts either toward (Engbert, R. & Kliegl, R. (2003). Microsaccades uncover the orientation of covert attention. Vision Research, 43, 1035-1045; Hafed, Z. M. & Clark, J. J. (2002). Microsaccades as an overt measure of covert attention shifts. Vision Research, 42(22), 2533-2545) or away from (Rolfs, M., Engbert, R., & Kliegl, R. (2004). Microsaccade orientation supports attentional enhancement opposite a peripheral cue: commentary on Tse, Sheinberg, and Logothetis (2003). Psychological Science, 15(10), 705-707) a peripheral cue under certain circumstances. Others (Horwitz, G. D. & Albright, T. D. (2003). Short-latency fixational saccades induced by luminance increments. Journal of Neurophysiology, 90(2), 1333-1339; Tse, P. U., Sheinberg, D. L., & Logothetis, N. K. (2002). Fixational eye movements are not affected by abrupt onsets that capture attention. Vision Research, 42, 1663-1669; Tse, P. U., Sheinberg, D. L., & Logothetis, N. K. (2004). The distribution of microsaccade directions need not reveal the location of attention. Psychological Science, 15(10), 708-710) found no change in the distribution of microsaccade directions as a function of where attention is allocated, although changes in the rate of microsaccades were observed in all of these studies in response to the onset of attentional reallocation. It is therefore possible that the distribution of microsaccade directions will change as a function of which figure is perceived to darken, or that changes in this distribution predict which figure will subsequently darken. We find no correlation between this distribution and which figure undergoes the effect, and therefore conclude that microsaccade directionality is not influenced by and does not influence which figure undergoes the effect. Moreover, the directions of microsaccades that occur immediately prior to a perceptual switch are not correlated with the perceived position of the figure that undergoes the effect. However, we do find that the rate of microsaccades decreases upon a perceptual switch, signifying an attentional shift coincident with the perceptual shift. We conclude that microsaccade directionality does not determine, predict, or cause which figure will subsequently be perceived to undergo an illusory brightness change. |
Massimo Turatto; Elena Betta Redundant visual signals boost saccade execution Journal Article In: Psychonomic Bulletin & Review, vol. 13, no. 5, pp. 928–932, 2006. @article{Turatto2006, The redundant signal effect (RSE) refers to the fact that human beings react more quickly to a pair of stimuli than to only one stimulus. In previous studies of the RSE in the oculomotor system, bimodal signals have been used as the goal of the saccade. In consistency with studies using manual response times (RTs), saccadic RTs have been shown to be shorter for redundant multimodal stimuli than for single unimodal stimuli. In the present experiments, we extended these findings by demonstrating an RSE in the saccadic system elicited only by unimodal visual stimuli. In addition, we found that shorter saccadic RTs were accompanied by an increased saccadic peak velocity. The present results are of relevance for neurophysiological models of saccade execution, since the boost of saccades was elicited by two visual transients (acting as a "go" signal) that were presented not at the goal of the saccade but at various other locations. |
Josef N. Geest; Gerardina C. Lagers-van Haselen; Maarten A. Frens Saccade adaptation in Williams-Beuren syndrome Journal Article In: Investigative Ophthalmology & Visual Science, vol. 47, no. 4, pp. 1464–1468, 2006. @article{Geest2006, PURPOSE: To investigate the capacity for rapid saccade adaptation in Williams-Beuren Syndrome (WBS), a genetic neurodevelopmental disorder, in which it has been observed that saccadic accuracy is severely reduced. METHODS: Saccade amplitude modification was elicited by backward steps (30% of target eccentricity) during the primary saccade in a classic saccade-adaptation paradigm. RESULTS: Patients with WBS showed a significant decrease in saccade amplitude. Furthermore, we observed that higher saccade accuracy before adaptation was related to more adaptation. CONCLUSIONS: The increased variability in motor performance does not abolish the ability for saccadic adaptation in subjects with WBS. Our results are congruent with the notion that part of the behavioral deficits observed in WBS may have a cerebellar origin. |
Wieske Zoest; Mieke Donk Saccadic target selection as a function of time Journal Article In: Spatial Vision, vol. 19, no. 1, pp. 61–76, 2006. @article{Zoest2006, Recent evidence indicates that stimulus-driven and goal-directed control of visual selection operate independently and in different time windows (van Zoest et al., 2004). The present study further investigates how eye movements are affected by stimulus-driven and goal-directed control. Observers were presented with search displays consisting of one target, multiple non-targets and one distractor element. The task of observers was to make a fast eye movement to a target immediately following the offset of a central fixation point, an event that either co-occurred with or soon followed the presentation of the search display. Distractor saliency and target-distractor similarity were independently manipulated. The results demonstrated that the effect of distractor saliency was transient and only present for the fastest eye movements, whereas the effect of target-distractor similarity was sustained and present in all but the fastest eye movements. The results support an independent timing account of visual selection. |
Robin Walker; Eugene McSorley The parallel programming of voluntary and reflexive saccades Journal Article In: Vision Research, vol. 46, no. 13, pp. 2082–2093, 2006. @article{Walker2006, A novel two-step paradigm was used to investigate the parallel programming of consecutive, stimulus-elicited ('reflexive') and endogenous ('voluntary') saccades. The mean latency of voluntary saccades, made following the first reflexive saccades in two-step conditions, was significantly reduced compared to that of voluntary saccades made in the single-step control trials. The latency of the first reflexive saccades was modulated by the requirement to make a second saccade: first saccade latency increased when a second voluntary saccade was required in the opposite direction to the first saccade, and decreased when a second saccade was required in the same direction as the first reflexive saccade. A second experiment confirmed the basic effect and also showed that a second reflexive saccade may be programmed in parallel with a first voluntary saccade. The results support the view that voluntary and reflexive saccades can be programmed in parallel on a common motor map. |
Robin Walker; Eugene Mcsorley; Patrick Haggard The control of saccade trajectories : Direction of curvature depends on prior knowledge of target location and saccade latency Journal Article In: Perception and Psychophysics, vol. 68, no. 1, pp. 129–138, 2006. @article{Walker2006a, Recent reports have shown that saccades can deviate either toward or away from distractors. However, the specific conditions responsible for the change in initial saccade direction are not known. One possibility, examined here, is that the direction of curvature (toward or away from distractors) reflects preparatory tuning of the oculomotor system when the location of the target and distractor are known in advance. This was investigated by examining saccade trajectories under predictable and unpredictable target conditions. In Experiment 1, the targets and the distractors appeared unpredictably, whereas in Experiment 2 an arrow cue presented at fixation indicated the location of the forthcoming target prior to stimulus onset. Saccades were made to targets on the horizontal, vertical, and principal oblique axis, and distractors appeared simultaneously at an adjacent location (a separation of +/- 45 degrees of visual angle). On average, saccade trajectories curved toward distractors when target locations were unpredictable and curved away from distractors when target locations were known in advance. There was no overall difference in mean saccade latencies between the two experiments. The magnitude of the distractor modulation of saccade trajectory (either toward or away from) was comparable across the different saccade directions (horizontal, vertical, and oblique). These results are interpreted in terms of the time course of competitive interactions operating in the neural structures involved in the suppression of distractors and the selection of a saccade target. A relatively slow mechanism that inhibits movements to distractors produces curvature away from the distractor. This mechanism has more time to operate when target location is predictable, increasing the likelihood that the saccade trajectory will deviate away from the distractor. |
Guy Wallis The temporal and spatial limits of compensation for fixational eye movements Journal Article In: Vision Research, vol. 46, no. 18, pp. 2848–2858, 2006. @article{Wallis2006, High-fidelity eye tracking is combined with a perceptual grouping task to provide insight into the likely mechanisms underlying the compensation of retinal image motion caused by movement of the eyes. The experiments describe the covert detection of minute temporal and spatial offsets incorporated into a test stimulus. Analysis of eye motion on individual trials indicates that the temporal offset sensitivity is actually due to motion of the eye inducing artificial spatial offsets in the briefly presented stimuli. The results have strong implications for two popular models of compensation for fixational eye movements, namely efference copy and image-based models. If an efference copy model is assumed, the results place constraints on the spatial accuracy and source of compensation. If an image-based model is assumed then limitations are placed on the integration time window over which motion estimates are calculated. |
Z. Wang; Louis F. Dell'Osso; Z. Zhang; R. John Leigh; Jonathan B. Jacobs Tenotomy does not affect saccadic velocities: Support for the "small-signal" gain hypothesis Journal Article In: Vision Research, vol. 46, no. 14, pp. 2259–2267, 2006. @article{Wang2006, We investigated the effects of four-muscle tenotomy on saccadic characteristics in infantile nystagmus syndrome (INS) and acquired pendular nystagmus (APN). Eye movements of 10 subjects with INS and one with APN were recorded using infrared reflection, magnetic search coil, or high-speed digital video. The expanded nystagmus acuity function (NAFX) quantified tenotomy-induced foveation changes in the INS. Saccadic characteristics and peak-to-peak nystagmus amplitudes were measured. Novel statistical tests were performed on the saccadic data. Six out of the 10 INS subjects showed no changes in saccadic duration, peak velocity, acceleration, or trajectory. In the other four, the differences were less than in peak-to-peak amplitudes (from 14.6% to 39.5%) and NAFX (from 22.2% to 162.4%). The APN subject also showed no changes despite a 50% decrease in peak-to-peak amplitude and a 34% increase in NAFX. The "small-signal" changes (peak-to-peak nystagmus amplitude and NAFX) were found to far exceed any "large-signal" changes (saccadic). Tenotomy successfully reduced INS and APN, enabling higher visual acuity without adversely affecting saccadic characteristics. These findings support the peripheral, small-signal gain reduction (via proprioceptive tension control) hypothesis. Current linear plant models, limited to normal steady-state muscle tension levels, cannot explain the effects of the tenotomy. |
Brian J. White; Dirk Kerzel; Karl R. Gegenfurtner The spatio-temporal tuning of the mechanisms in the control of saccadic eye movements Journal Article In: Vision Research, vol. 46, no. 22, pp. 3886–3897, 2006. @article{White2006, We compared the spatio-temporal tuning of perception to the mechanisms that drive saccadic eye movements. Detection thresholds were measured for Gabor-targets presented left or right of fixation (4 or 8 deg eccentricity), at one of four spatial frequencies (1, 2, 4 or 8 cpd) oscillating at one of three temporal frequencies (1, 8 or 16 Hz). We then measured saccade latency to each target presented at various multiples of detection threshold. Consistent with previous research, latency decreased as a function of contrast. However, at equal detection performance, we found no systematic difference in saccadic latency and no difference in average oculometric performance (% correct saccade direction) across the different target spatio-temporal frequencies. Furthermore, position error remained fairly constant across all conditions. The results are consistent with the idea that the spatio-temporal signals used for perception are the same as those used by the mechanisms driving saccadic eye movements. |
F. Møller; M. L. Laursen; A. K. Sjølie Fixation topography in normal test persons Journal Article In: Graefe's Archive for Clinical and Experimental Ophthalmology, vol. 244, pp. 577–582, 2006. @article{Moeller2006, BACKGROUND: The eye is moved so that the object of interest falls on the central fovea, where the spatial resolution is highest. In the present study we quantified eye movements of normal test persons during steady fixation and characterized the fixation using a 3D fixation plot (X horizontal eye position, Y vertical eye position, Z time in each eye position). METHOD: Fixation eye movements were quantified binocularly in ten normal test persons during a 40-s fixation task using an infrared recording technique. RESULTS: The fixation plot was characterized by a single preferred fixation locus in 17 eyes. One eye had two distinctly separated preferred fixation locations and in two eyes the configuration of fixation plot was flat with no single identifiable locus of fixation. The fixation plots were elliptical along the horizontal meridian in 9 eyes, elliptical along the vertical meridian in 8 eyes, and round in 3 eyes. The fixation area (RAF95) ranged between 1418 and 14182 arcmin(2), and a significant positive correlation was found between RAF95 and the mean microsaccadic amplitude (p<0.001). CONCLUSION: The fixation plots are often characterized by a single preferred fixation locus but may also be almost flat with no identifiable location of fixation. The individual fixations patterns resembles the cone density contour plots as found in histological studies, and it may be speculated, that the shape of the fixation plot is determined by the cone density topography. |
F. Møller; M. L. Laursen; A. K. Sjølie The contribution of microsaccades and drifts in the maintenance of binocular steady fixation Journal Article In: Graefe's Archive for Clinical and Experimental Ophthalmology, vol. 244, no. 4, pp. 465–471, 2006. @article{Moeller2006a, BACKGROUND: The eye performs three types of eye movements during fixation: fast microsaccades are interrupted by slow drift movements, and tremor is superimposed on the drifts. The contribution of the microsaccades and drifts in maintaining fixation has been discussed since the late 1950s. Initially, microsaccades were thought to correct the misalignment from the optimal fixation locus induced by the drift movements, a theory still postulated in more recent work. The present study aimed to uncover to what extent each fixation movement contributes to maintain steady binocular fixation. METHOD: Binocular fixation during a 40-s fixation task was recorded using an infrared recording technique for ten normal test persons. Start and end point of each microsaccade and drift were superimposed on a fixation map, and the distance to the preferred retinal location of fixation (PRL) was measured. RESULTS: It was found that 32.6% of the microsaccades corrected the previous drift movement towards the PRL, whereas 53.1% of the drifts corrected the endpoint of the previous microsaccade towards the PRL. The overall mean post-microsaccadic and mean post-drift distance to the PRL for the ten normal test persons were 0.46 degrees and 0.41 degrees , respectively; the difference was not statistically significant. Interindividually, the mean post-microsaccadic distance to the PRL ranged between 0.21 degrees and 0.91 degrees and the mean post-drift distance to the PRL ranged between 0.20 degrees and 0.72 degrees . CONCLUSION: Neither the endpoints of the microsaccades nor the drifts bring the visual line to coincide with the centre of the PRL. Consequently, it must be the eye movements performed during the drifts ("slow control") that keep the visual line in the centre of the foveola. |
Leigh A. Mrotek; Martha Flanders; John F. Soechting Oculomotor responses to gradual changes in target direction Journal Article In: Experimental Brain Research, vol. 172, no. 2, pp. 175–192, 2006. @article{Mrotek2006, Smooth pursuit tracking of targets moving linearly (in one dimension) is well characterized by a model where retinal image motion drives eye acceleration. However, previous findings suggest that this model cannot be simply extended to two-dimensional (2D) tracking. To examine 2D pursuit, in the present study, human subjects tracked a target that moved linearly and then followed the arc of a circle. The subjects' gaze angular velocity accurately matched target angular velocity, but the direction of smooth pursuit always lagged behind the current target direction. Pursuit speed slowly declined after the onset of the curve (for about 500 ms), even though the target speed was constant. In a second experiment, brief perturbations were presented immediately prior to the beginning of the change in direction. The subjects' responses to these perturbations consisted of two components: (1) a response specific to the parameters of the perturbation and (2) a nonspecific response that always consisted of a transient decrease in gaze velocity. With the exception of this nonspecific response, pursuit behavior in response to the gradual changes in direction and to the perturbations could be explained by using retinal slip (image velocity) as the input signal. The retinal slip was parallel and perpendicular to the instantaneous direction of pursuit ultimately resulted in changes in gaze velocity (via gaze acceleration). Perhaps due to the subjects' expectations that the target will curve, the sensitivity to the image motion in the direction of pursuit was not as strong as the sensitivity to image motion perpendicular to gaze velocity. |
Lauri Nummenmaa; Jari K. Hietanen Gaze distractors influence saccadic curvature: Evidence for the role of the oculomotor system in gaze-cued orienting Journal Article In: Vision Research, vol. 46, no. 21, pp. 3674–3680, 2006. @article{Nummenmaa2006, We examined the role of the oculomotor system in gaze-triggered orienting of attention by measuring whether perceiving of task-irrelevant gaze distractors and peripheral spatial distractors influence saccadic curvature similarly. Participants performed reflexive, vertical saccades to designated target areas while their eye movements were recorded. Schematic faces with averted gaze or peripheral boxes were presented before or simultaneously (-100 ms/0 ms SOAs) with the imperative signal. Gaze distractors caused the saccades to curve away from the distractor direction at both SOAs and peripheral distractors only at the 0-ms SOA. The results imply that gaze-cued shifts of visual attention involve both cortical attention orienting systems and subcortical oculomotor systems. |
Susana Martinez-Conde; Stephen L. Macknik; Xoana G. Troncoso; Thomas A. Dyar Microsaccades counteract visual fading during fixation Journal Article In: Neuron, vol. 49, no. 2, pp. 297–305, 2006. @article{MartinezConde2006, Our eyes move continually, even while we fixate our gaze on an object. If fixational eye movements are counteracted, our perception of stationary objects fades completely, due to neural adaptation. Some studies have suggested that fixational microsaccades refresh retinal images, thereby preventing adaptation and fading. However, other studies disagree, and so the role of microsaccades remains unclear. Here, we correlate visibility during fixation to the occurrence of microsaccades. We asked subjects to indicate when Troxler fading of a peripheral target occurs, while simultaneously recording their eye movements with high precision. We found that before a fading period, the probability, rate, and magnitude of microsaccades decreased. Before transitions toward visibility, the probability, rate, and magnitude of microsaccades increased. These results reveal a direct link between suppression of microsaccades and fading and suggest a causal relationship between microsaccade production and target visibility during fixation. |
Eugene McSorley Time course of oculomotor inhibition revealed by saccade trajectory modulation Journal Article In: Journal of Neurophysiology, vol. 96, no. 3, pp. 1420–1424, 2006. @article{McSorley2006, Selecting a stimulus as the target for a goal-directed movement involves inhibiting other competing possible responses. Both target and distractor stimuli activate populations of neurons in topographic oculomotor maps such as the superior colliculus. Local inhibitory interconnections between these populations ensure only one saccade target is selected. Suppress-ing saccades to distractors may additionally involve inhibiting corre-sponding map regions to bias the local competition. Behavioral evidence of these inhibitory processes comes from the effects of distractors on oculomotor and manual trajectories. Individual saccades may initially deviate either toward or away from a distractor, but the source of this variability has not been investigated. Here we investi-gate the relation between distractor-related deviation of trajectory and saccade latency. Targets were presented with, or without, distractors, and the deviation of saccade trajectories arising from the presence of distractors was measured. A fixation gap paradigm was used to manipulate latency independently of the influence of competing dis-tractors. Shorter-latency saccades deviated toward distractors and longer-latency saccades deviated away from distractors. The transition between deviation toward or away from distractors occurred at a saccade latency of around 200 ms. This shows that the time course of the inhibitory process involved in distractor related suppression is relatively slow. |
Ibrahim Dahlstrom-Hakki; Alexander Pollatsek Limits on integrating motion information across saccades Journal Article In: Perception and Psychophysics, vol. 68, no. 1, pp. 43–54, 2006. @article{DahlstromHakki2006, In two experiments, we investigated whether people could detect changes in the rotary motion of a cube. A rendering of a cube rotating at a constant angular velocity was presented on a video monitor and, at a key point in the trial, a cross was presented to one side of the cube as a cue for a saccade. On some trials, a change in the rotation occurred either about 100 msec before the saccade or during the saccade; on other trials, there was no change. The change consisted of moving the cube to a new position in the "rotation sequence," after which it continued to rotate at the same angular velocity as before. There was also a control on all trials to ensure that change detection was not due to the detection of low-level motion. Although detection of the change was well above chance when it occurred during the fixation, it was at chance when it occurred during the saccade, except in the case of one participant (who was in both experiments). This chance performance also occurred in Experiment 2 for (1) a slower rotation speed and (2) an axis of rotation that made the rotation planar. The participant who had above chance performance (and as good as that when the change occurred during a fixation) reported using a "strategy" that did not track the path of the cube. It thus appears that there is no natural way in which the visual system tracks this rotary motion, and that detection of change requires some sort of recoding. This finding raises the question of whether good performance in other, apparently similar, motion-detection tasks is a result of similar recoding. |
Elena Betta; Massimo Turatto Are you ready? I can tell by looking at your microsaccades Journal Article In: NeuroReport, vol. 17, no. 10, pp. 1001–1004, 2006. @article{Betta2006, The direction of microsaccades has been shown to be biased by the allocation of spatial attention. Here, we investigated whether the cognitive processes involved in preparing to respond to an upcoming target can modulate the microsaccadic response. Specifically, we found that optimal manual response preparation, reflected by faster response times, was associated with a reduction in the absolute frequency of microsaccades. The present results are consistent with previous studies suggesting a relationship between oculomotor activity and different sorts of motor responses. Our findings, however, surprisingly demonstrate that the effect of preparation and stimulus expectation extends to an automatic and unconscious oculomotor activity such as microsaccade execution. |
2005 |
Christian Bellebaum; Irene Daum; B. Koch; M. Schwarz; Klaus-Peter Hoffmann The role of the human thalamus in processing corollary discharge Journal Article In: Brain, vol. 128, no. 5, pp. 1139–1154, 2005. @article{Bellebaum2005, Corollary discharge signals play an important role in monitoring self-generated movements to guarantee spatial constancy. Recent work in macaques suggests that the thalamus conveys corollary discharge information of upcoming saccades passing from the superior colliculus to the frontal eye field. The present study aimed to investigate the involvement of the thalamus in humans by assessing the effect of thalamic lesions on the processing of corollary discharge information. Thirteen patients with selective thalamic lesions and 13 healthy age-matched control subjects performed a saccadic double-step task in which retino-spatial dissonance was induced, i.e. the retinal vector of the second target and the movement vector of the second saccade were different. Thus, the subjects could not rely on retinal information alone, but had to use corollary discharge information to correctly perform the second saccade. The amplitudes of first and second saccades were significantly smaller in patients than in controls. Five thalamic lesion patients showed unilateral deficits in using corollary discharge information, as revealed by asymmetries compared with the other patients and controls. Three patients with lateral thalamic lesions including the ventrolateral nucleus (VL) were impaired contralaterally to the side of damage and one patient with a lesion in the mediodorsal thalamus (MD) was impaired ipsilaterally to the lesion. The largest asymmetry was found in a patient with a bilateral thalamic lesion. The results provide evidence for a thalamic involvement in the processing of corollary discharge information in humans, with a potential role of both the VL and MD nuclei |
Christian Bellebaum; Klaus-Peter Hoffmann; Irene Daum Post-saccadic updating of visual space in the posterior parietal cortex in humans Journal Article In: Behavioural Brain Research, vol. 163, no. 2, pp. 194–203, 2005. @article{Bellebaum2005a, Updating of visual space takes place in the posterior parietal cortex to guarantee spatial constancy across eye movements. However, the timing of updating with respect to saccadic eye movements remains a matter of debate. In the present study, event-related potentials (ERPs) were recorded in 15 volunteers during a saccadic double-step task to elucidate the time course of the updating process. In the experimental condition updating of visual space was required, because both saccade targets had already disappeared before the first saccade was executed. A similar task without updating requirements served as control condition. ERP analysis revealed a significantly larger slow positive wave in the retino-spatial dissonance condition compared to the control condition, starting between 150 and 200 ms after first saccade onset. Source analysis showed an asymmetry with respect to the direction of the first saccade. Whereas the source was restricted to the right PPC in trials with leftward first saccades, left and right PPC were involved in rightward trials. The results of the present study suggest that updating of visual space in a saccadic double-step task occurs not earlier than 150 ms after the onset of the first saccade. We conclude that extraretinal information about the first saccade is integrated with motor information about the second saccade in the inter-saccade interval. |
F. Møller; M. L. Laursen; A. K. Sjølie Binocular fixation topography in patients with diabetic macular oedema: Possible implications for photocoagulation therapy Journal Article In: Graefe's Archive for Clinical and Experimental Ophthalmology, vol. 243, no. 9, pp. 903–910, 2005. @article{Moeller2005, During retinal photocoagulation for diabetic maculopathy, there is a potential risk of foveal burns, and laser scars may later enlarge to be sight-threatening when involving retinal areas previously used during fixation. Since the retinal area used during binocular steady fixation has been found to vary considerably in the normal test person and central fixation may be even further compromised in patients with diabetic maculopathy, the sight-threatening side effects could possibly be reduced by taking into account the fixation area individually. However, no study has described and quantified the retinal area of fixation binocularly in patients with clinically significant macular oedema (CSME). |
Anna Montagnini; Leonardo Chelazzi The urgency to look: Prompt saccades to the benefit of perception Journal Article In: Vision Research, vol. 45, no. 27, pp. 3391–3401, 2005. @article{Montagnini2005, Researchers have shown that the promptness to initiate a saccade is modulated by countless factors pertaining to the visual context and the task. However, experiments on saccadic eye movements are usually designed in such a way that oculomotor performance is dissociated from the natural role of saccades, namely that of making crucial perceptual information rapidly available for high-resolution, foveal analysis. Here, we demonstrate that the requirement to perform a difficult perceptual judgment at the saccade landing location can reduce saccadic latency (by >15%) and increase saccadic peak velocity. Importantly, the effect cannot be explained in terms of arousal, as latency changes are specific to the location where the perceptual judgement is required. These results indicate that mechanisms for voluntary saccade initiation are under the powerful indirect control of perceptual goals. |
Shay Moshel; Jinrong Liang; Avi Caspi; Ralf Engbert; Reinhold Kliegl; Shlomo Havlin; Ari Z. Zivotofsky Phase-synchronization decay of fixational eye movements Journal Article In: Annals of the New York Academy of Sciences, vol. 1039, pp. 484–488, 2005. @article{Moshel2005, In nonstationary noisy systems the traditional cross-correlation method may not appropriately detect all cases of interdependencies between coupled systems. The phase-synchronization method was previously found use-ful in detecting synchronization in several systems. We here applied the phase-synchronization decay to study the synchronization between six combinations of binocular fixational eye movement components. We found that only two compo-nents were synchronized: the right and left horizontal with each other and the right and left vertical. Furthermore, the vertical-vertical components were much more synchronized than the horizontal. |
Nadia Alahyane; Denis Pélisson Long-lasting modifications of saccadic eye movements following adaptation induced in the double-step target paradigm Journal Article In: Learning and Memory, vol. 12, no. 4, pp. 433–443, 2005. @article{Alahyane2005, The adaptation of saccadic eye movements to environmental changes occurring throughout life is a good model of motor learning and motor memory. Numerous studies have analyzed the behavioral properties and neural substrate of oculomotor learning in short-term saccadic adaptation protocols, but to our knowledge, none have tested the persistence of the oculomotor memory. In the present study, the double-step target protocol was used in five human subjects to adaptively decrease the amplitude of reactive saccades triggered by a horizontally-stepping visual target. We tested the amplitude of visually guided saccades just before and at different times (up to 19 days) after the adaptation session. The results revealed that immediately after the adaptation session, saccade amplitude was significantly reduced by 22% on average. Although progressively recovering over days, this change in saccade gain was still statistically significant on days 1 and 5, with an average retention rate of 36% and 19%, respectively. On day 11, saccade amplitude no longer differed from the pre-adaptation value. Adaptation was more effective and more resistant to recovery for leftward saccades than for rightward ones. Lastly, modifications of saccade gain related to adaptation were accompanied by a decrease of both saccade duration and peak velocity. A control experiment indicated that all these findings were specifically related to the adaptation protocol, and further revealed that no change in the main sequence relationships could be specifically related to adaptation. We conclude that in humans, the modifications of saccade amplitude that quickly develop during a double-step target adaptation protocol can remain in memory for a much longer period of time, reflecting enduring plastic changes in the brain. |
Nadia Alahyane; Denis Pélisson Retention of saccadic adaptation in humans Journal Article In: Annals of the New York Academy of Sciences, vol. 1039, pp. 558–562, 2005. @article{Alahyane2005a, In the present study, we tested in human subjects the persistence of the oculomotor changes resulting from saccadic adaptation up to 19 days after exposure to the double step target protocol. The main results indicate that the reduction of saccade gain related to the adaptation session (mean gain change of 5 subjects = 22 +/- 4.7%) was partially but significantly retained after 1 day and 5 days (mean amount of retention = 36 +/- 17% and 19.7 +/- 13.3%, respectively) but was no longer significant at day 11 and 19. Unexpectedly, gain changes were larger for leftward than for rightward saccades. No change in saccade dynamics was observed. These data suggest that in humans, adaptive mechanisms induce long lasting changes in visually-guided saccade amplitude, probably reflecting plastic changes in the brain. |
Dirk Kerzel; Nathalie E. Ziegler Visual short-term memory during smooth pursuit eye movements Journal Article In: Journal of Experimental Psychology: Human Perception and Performance, vol. 31, no. 2, pp. 354–372, 2005. @article{Kerzel2005a, Large-scale whole genome association studies are increasingly common, due in large part to recent advances in genotyping technology. With this change in paradigm for genetic studies of complex diseases, it is vital to develop valid, powerful, and efficient statistical tools and approaches to evaluate such data. Despite a dramatic drop in genotyping costs, it is still expensive to genotype thousands of individuals for hundreds of thousands single nucleotide polymorphisms (SNPs) for large-scale whole genome association studies. A multi-stage (or two-stage) design has been a promising alternative: in the first stage, only a fraction of samples are genotyped and tested using a dense set of SNPs, and only a small subset of markers that show moderate associations with the disease will be genotyped in later stages. Multi-stage designs have also been used in candidate gene association studies, usually in regions that have shown strong signals by linkage studies. To decide which set of SNPs to be genotyped in the next stage, a common practice is to utilize a simple test (such as a chi2 test for case-control data) and a liberal significance level without corrections for multiple testing, to ensure that no true signals will be filtered out. In this paper, I have developed a novel SNP selection procedure within the framework of multi-stage designs. Based on data from stage 1, the method explicitly explores correlations (linkage disequilibrium) among SNPs and their possible interactions in determining the disease phenotype. Comparing with a regular multi-stage design, the approach can select a much reduced set of SNPs with high discriminative power for later stages. Therefore, not only does it reduce the genotyping cost in later stages, it also increases the statistical power by reducing the number of tests. Combined analysis is proposed to further improve power, and the theoretical significance level of the combined statistic is derived. Extensive simulations have been performed, and results have shown that the procedure can reduce the number of SNPs required in later stages, with improved power to detect associations. The procedure has also been applied to a real data set from a genome-wide association study of the sporadic amyotrophic lateral sclerosis (ALS) disease, and an interesting set of candidate SNPs has been identified. |
Jochen Laubrock; Ralf Engbert; Reinhold Kliegl Microsaccade dynamics during covert attention Journal Article In: Vision Research, vol. 45, no. 6, pp. 721–730, 2005. @article{Laubrock2005, We compared effects of covert spatial-attention shifts induced with exogenous or endogenous cues on microsaccade rate and direction. Separate and dissociated effects were obtained in rate and direction measures. Display changes caused microsaccade rate inhibition, followed by sustained rate enhancement. Effects on microsaccade direction were differentially tied to cue class (exogenous vs. endogenous) and type (neutral vs. directional). For endogenous cues, direction effects were weak and occurred late. Exogenous cues caused a fast direction bias towards the cue (i.e., early automatic triggering of saccade programs), followed by a shift in the opposite direction (i.e, controlled inhibition of cue-directed saccades, leading to a 'leakage' of microsaccades in the opposite direction). |
John F. Soechting; Leigh A. Mrotek; Martha Flanders Smooth pursuit tracking of an abrupt change in target direction: Vector superposition of discrete responses Journal Article In: Experimental Brain Research, vol. 160, no. 2, pp. 245–258, 2005. @article{Soechting2005, The directional control of smooth pursuit eye movements was studied by presenting human subjects with targets that moved in a straight line at a constant speed and then changed direction abruptly and unpredictably. To minimize the probability of saccadic responses in the interval following the target's change in direction, target position was offset so as to eliminate position error after the reaction time. Smooth pursuit speed declined at a latency of 90 ms, whereas the direction of smooth pursuit began to change later (130 ms). The amplitude of the offset in target position did not affect the subsequent smooth pursuit response. In other experiments, the target's speed or acceleration was changed abruptly at the time of the change in direction. Step changes in speed elicited short-latency responses in smooth pursuit tracking but step changes in acceleration did not. In all instances, the earliest component of the response did not depend on the parameters of the stimulus. The data were fit with a model in which smooth pursuit resulted from the vector addition of two components, one representing a response to the arrest of the initial target motion and the other the response to the onset of target motion in the new direction. This model gave an excellent fit but further analysis revealed nonlinear interactions between the two vector components. These interactions represented directional anisotropies both in terms of the initial tracking direction (which was either vertical or 45 degrees ) and in terms of the cardinal directions (vertical and horizontal). |
Jan L. Souman; Ignace T. C. Hooge; Alexander H. Wertheim Perceived motion direction during smooth pursuit eye movements Journal Article In: Experimental Brain Research, vol. 164, no. 3, pp. 376–386, 2005. @article{Souman2005, Although many studies have been devoted to motion perception during smooth pursuit eye movements, relatively little attention has been paid to the question of whether the compensation for the effects of these eye movements is the same across different stimulus directions. The few studies that have addressed this issue provide conflicting conclusions. We measured the perceived motion direction of a stimulus dot during horizontal ocular pursuit for stimulus directions spanning the entire range of 360 degrees. The stimulus moved at either 3 or 8 degrees/s. Constancy of the degree of compensation was assessed by fitting the classical linear model of motion perception during pursuit. According to this model, the perceived velocity is the result of adding an eye movement signal that estimates the eye velocity to the retinal signal that estimates the retinal image velocity for a given stimulus object. The perceived direction depends on the gain ratio of the two signals, which is assumed to be constant across stimulus directions. The model provided a good fit to the data, suggesting that compensation is indeed constant across stimulus direction. Moreover, the gain ratio was lower for the higher stimulus speed, explaining differences in results in the literature. |
Jan L. Souman; Ignace T. C. Hooge; Alexander H. Wertheim Vertical object motion during horizontal ocular pursuit: Compensation for eye movements increases with presentation duration Journal Article In: Vision Research, vol. 45, no. 7, pp. 845–853, 2005. @article{Souman2005a, Smooth pursuit eye movements change the retinal image motion of objects in the visual field. To enable an observer to perceive the motion of these objects veridically, the visual system has to compensate for the effects of the eye movements. The occurrence of the Filehne-illusion (illusory motion of a stationary object during smooth pursuit) shows that this compensation is not always perfect. The amplitude of the illusion appears to decrease with increasing presentation durations of the stationary object. In this study we investigated whether presentation duration has the same effect when an observer views a vertically moving object during horizontal pursuit. In this case, the pursuit eye movements cause the perceived motion path to be oblique instead of vertical; this error in perceived motion direction should decrease with higher presentation durations. In Experiment 1, we found that the error in perceived motion direction indeed decreased with increasing presentation duration, especially for higher pursuit velocities. The results of Experiment 2 showed that the error in perceived motion direction did not depend on the moment during pursuit at which the stimulus was presented, suggesting that the degree of compensation for eye movements is constant throughout pursuit. The results suggest that longer presentation durations cause the eye movement signal that is used by the visual system to increase more than the retinal signal. |
Miriam Spering; Dirk Kerzel; Doris I. Braun; Michael J. Hawken; Karl R. Gegenfurtner Effects of contrast on smooth pursuit eye movements Journal Article In: Journal of Vision, vol. 5, pp. 455–465, 2005. @article{Spering2005, It is well known that moving stimuli can appear to move more slowly when contrast is reduced (P. Thompson, 1982). Here we address the question whether changes in stimulus contrast also affect smooth pursuit eye movements. Subjects were asked to smoothly track a moving Gabor patch. Targets varied in velocity (1, 8, and 15 deg/s), spatial frequency (0.1, 1, 4, and 8 c/deg), and contrast, ranging from just below individual thresholds to maximum contrast. Results show that smooth pursuit eye velocity gain rose significantly with increasing contrast. Below a contrast level of two to three times threshold, pursuit gain, acceleration, latency, and positional accuracy were severely impaired. Therefore, the smooth pursuit motor response shows the same kind of slowing at low contrast that was demonstrated in previous studies on perception |
Leo Trottier; Jay Pratt Visual processing of targets can reduce saccadic latencies Journal Article In: Vision Research, vol. 45, no. 11, pp. 1349–1354, 2005. @article{Trottier2005, Normal human saccadic reaction times (SRTs) have been thought to be approximately 200 ms. The present study, using an experimental method that takes advantage of what the saccade system has evolved to do (by instructing subjects to rapidly acquire detailed visual information from the environment), shows that human SRTs are actually on the order of 150 ms. Moreover, when combined with the sensory-based "gap" effect (removal of gaze fixation object prior to target presentation), this method yielded extremely low SRTs. These findings imply that previous approximations of human SRTs may have been too conservative, and that the group of saccades often classified as "express" may instead represent the norm. |
Loes C. J. Dam; Raymond Ee The role of (micro)saccades and blinks in perceptual bi-stability from slant rivalry Journal Article In: Vision Research, vol. 45, no. 18, pp. 2417–2435, 2005. @article{Dam2005, We exposed the visual system to an ambiguous 3D slant rivalry stimulus consisting of a grid for which monocular (perspective) and binocular (disparity) cues independently specified a slant about a horizontal axis. When those cues specified similar slants, observers perceived a single slant. When the difference between the specified slants was large, observers alternatively perceived a perspective- or a disparity-dominated slant. Eye movement measurements revealed that there was no positive correlation between a perceptual flip and both saccades (microsaccades as well as larger saccades) and blinks that occurred prior to a perceptual flip. We also found that changes in horizontal vergence were not responsible for perceptual flips. Thus, eye movements were not essential to flip from one percept to the other. After the moment of a perceptual flip the occurrence probabilities of both saccades and blinks were reduced. The reduced probability of saccades mainly occurred for larger voluntary saccades, rather than for involuntary microsaccades. We suggest that the reduced probability of voluntary saccades reflects a reset of saccade planning. |
Brian J. White; Karl R. Gegenfurtner; Dirk Kerzel Effects of structured nontarget stimuli on saccadic latency Journal Article In: Journal of Neurophysiology, vol. 93, no. 6, pp. 3214–3223, 2005. @article{White2005, It has been suggested that the remote distractor effect is the result of nontarget stimulation of a central region representing a collicular fixation zone near the time of target onset. The distributed network of the cells responsible for this effect is believed to extend over a large area, responding to distractors < or =10 deg in the periphery. Several studies also implicate the superior colliculus as the substrate behind an inhibited saccadic response arising from a display change. We investigated this further by using a patch of pink noise of various sizes as a nontarget stimulus. We show that the onset of a small patch (2.3 x 2.3 deg) of centrally displayed pink noise can produce a significant increase in saccadic latency to a simultaneously presented peripheral Gabor target. In contrast, a large patch (36 x 36 deg) of pink noise did not increase latency despite the fact that it also stimulated the region representing the fixation zone. Furthermore, only the large patch of noise facilitated latency when presented before target onset. We also examined the effect of patch sizes between these two extremes and found a steady decrease in latency as patch size increased. This confirms that nontarget stimulation of the region representing the fixation zone near the time of target onset is not in itself sufficient to produce the increase in latency typically found with remote distractors. The results are consistent with the idea that only a spatially confined object leads to a discharge of collicular fixation neurons. |
Valérie Gaveau; Nadia Alahyane; Roméo Salemme; Michel Desmurget Self-generated saccades do not modify the gain of adapted reactive saccades Journal Article In: Experimental Brain Research, vol. 162, no. 4, pp. 526–531, 2005. @article{Gaveau2005, The gain of reactive saccades was manipulated in 17 subjects using a target-jump paradigm. Following adaptation three sub-groups were formed: (1) rest 15 min in the dark, eyes closed; (2) perform self-generated saccades for 15 min; (3) perform reactive saccades for 15 min. The series of saccades were the same in groups 2 and 3 (amplitude, sequence), except that group 3 performed fewer saccades (same number as the lowest number of saccades performed by one subject in group 2). Neither the rest period nor the series of self-generated saccades affected the adapted gain. The series of reactive saccades generated, by contrast, quick de-adaptation. These results support the conclusion that the gain of self-generated and reactive saccades is independently controlled. |
Anne Lise V. Holahan; Gillian A. O'Driscoll Antisaccade and smooth pursuit performance in positive- and negative-symptom schizotypy Journal Article In: Schizophrenia Research, vol. 76, no. 1, pp. 43–54, 2005. @article{Holahan2005, Schizophrenic patients have well-documented abnormalities in smooth pursuit eye movements and antisaccade performance. In populations at risk for schizophrenia, smooth pursuit abnormalities are also well documented. Antisaccade deficits have been replicated in high-risk populations as well, but the findings are more variable and the reasons for the variability are not clear. Some evidence suggests that antisaccade deficits increase in high-risk populations in relation to the presence of positive symptoms. Whether antisaccade deficits increase in relation to negative symptoms in high-risk populations is relatively uninvestigated. We evaluated antisaccade and pursuit performance in "psychometric schizotypes" who had elevated scores on either the Perceptual Aberration Scale (PerAb; i.e., positive symptoms) or the Physical Anhedonia Scale (PhysAnh; i.e., negative symptoms) but not both, and in normal controls. We used the standard version of the antisaccade task, for which results in positive-symptom schizotypes have previously been reported, and investigated performance on a gap and overlap version. We replicated the finding that a significantly larger percentage of positive-symptom schizotypes than controls have elevated antisaccade error rates on the standard antisaccade task (P=0.03); the percentage of negative-symptom schizotypes with elevated antisaccade error rates did not differ from that of control subjects. Neither schizotypal group was impaired on the gap or overlap versions of the task. On the pursuit task, a higher percentage of positive- and negative-symptom schizotypes were classified as having deviant performance than control subjects (both Ps<0.04). These findings suggest that antisaccade deficits may be better at identifying high-risk subjects with positive symptoms. Pursuit deficits identified both positive- and negative-symptom schizotypes, but was better at identifying the latter. |
Samuel B. Hutton; D. Tegally The effects of dividing attention on smooth pursuit eye tracking Journal Article In: Experimental Brain Research, vol. 163, no. 3, pp. 306–313, 2005. @article{Hutton2005, Attentional processes have traditionally been closely linked to the production of saccadic eye movements, but their role in the control of smooth pursuit eye movements remains unclear. In two experiments we used dual task paradigms to vary the attentional resources available for pursuit eye tracking. In both experiments we found that attentionally demanding secondary tasks impaired smooth pursuit performance, resulting in decreased velocity and increased position error. These findings suggest that attention is important for the maintenance of accurate smooth pursuit, and do not support the hypothesis that pursuit is a relatively automatic function that proceeds optimally in the absence of attentional control. These results add weight to the suggestion that a similar functional architecture underlies both pursuit and saccadic eye movements. |
Jin Rong Liang; Shay Moshel; Ari Z. Zivotofsky; Avi Caspi; Ralf Engbert; Reinhold Kliegl; Shlomo Havlin Scaling of horizontal and vertical fixational eye movements Journal Article In: Physical Review E, vol. 71, no. 3, pp. 1–6, 2005. @article{Liang2005, Eye movements during fixation of a stationary target prevent the adaptation of the photoreceptors to continuous illumination and inhibit fading of the image. These random, involuntary, small, movements are restricted at long time scales so as to keep the target at the center of the field of view. Here we use the Detrended Fluctuation Analysis (DFA) in order to study the properties of fixational eye movements at different time scales. Results show different scaling behavior between horizontal and vertical movements. When the small ballistics movements, i.e. micro-saccades, are removed, the scaling exponents in both directions become similar. Our findings suggest that micro-saccades enhance the persistence at short time scales mostly in the horizontal component and much less in the vertical component. This difference may be due to the need of continuously moving the eyes in the horizontal plane, in order to match the stereoscopic image for different viewing distance. |
Eugene McSorley; Patrick Haggard; Robin Walker Spatial and temporal aspects of oculomotor inhibition as revealed by saccade trajectories Journal Article In: Vision Research, vol. 45, no. 19, pp. 2492–2499, 2005. @article{McSorley2005, The spatial and temporal effect of distractor related inhibition on stimulus elicited (reflexive) and goal driven (voluntary) saccades, was examined using saccade trajectory deviations as a measure. Subjects made voluntary and reflexive saccades to a target location on the vertical midline, while the distance of a distractor from the target was systematically manipulated. The trajectory curvature of both voluntary and reflexive saccades was found to be subject to individual differences. Saccade curvature was found to decrease monotonically with increasing distractor distance from target for some subjects while for others no reduction in curvature or even an increase was found. These results could not be explained by latency differences or landing position effects. The different patterns of distractor effects on saccade trajectories suggest the additional influence of a non-spatial inhibitory mechanism. |
2004 |
Casimir J. H. Ludwig; Iain D. Gilchrist; Eugene McSorley The influence of spatial frequency and contrast on saccade latencies Journal Article In: Vision Research, vol. 44, no. 22, pp. 2597–2604, 2004. @article{Ludwig2004, We characterised the impact of spatial frequency and contrast on saccade latencies to single Gabor patches. Saccade latencies decreased as a function of contrast, and increased with spatial frequency. The observed latency variations are qualitatively similar to those observed for manual reaction times. For single target detection, our findings highlight the similarity in the visual processes that support both saccadic and manual responses. |
Eugene McSorley; Patrick Haggard; Robin Walker Distractor modulation of saccade trajectories: Spatial separation and symmetry affects Journal Article In: Experimental Brain Research, vol. 155, no. 3, pp. 320–333, 2004. @article{McSorley2004, The trajectories of saccadic eye movements can be modulated by the presence of a competing visual distractor. In the present study the trajectories of vertical saccades curved away from a single visual distractor presented in one visual field, but tended to be straight when two distractors were presented at mirror symmetric locations in both visual fields. The spatial nature of the mirror distractor effect was examined by presenting a second distractor at mirror and non-mirror locations. Saccade trajectories also tended to be straight with both mirror and non-mirror symmetrical distractors. The relationship between the distractor location and saccade curvature was examined in a third experiment by manipulating the distractor-to-target spatial separation. Although there was a tendency for greater curvature when the distractor was presented in the same hemifield as the target there was no clear relationship between curvature and distractor location. The results show that the distractor modulation of saccade trajectory is not highly spatially specific and that it can be balanced by a second bilateral distractor in the opposite visual field. The results are interpreted in terms of a model in which the initial saccade direction and curvature back towards the saccade goal are controlled by separate processes. Initial saccade direction is modulated by the inhibition of distractor locations within a 'motor map' specifying saccade direction. Curvature back towards the saccade goal may be attributed to a feedback system, with a separate representation of the visual target location, that enables an on-line correction of the saccade during mid-flight. |
Wieske Zoest; Mieke Donk; Jan Theeuwes The role of stimulus-driven and goal-driven control in saccadic visual selection Journal Article In: Journal of Experimental Psychology: Human Perception and Performance, vol. 30, no. 4, pp. 746–759, 2004. @article{Zoest2004, Four experiments were conducted to investigate the role of stimulus-driven and goal-driven control in saccadic eye movements. Participants were required to make a speeded saccade toward a predefined target presented concurrently with multiple nontargets and possibly 1 distractor. Target and distractor were either equally salient (Experiments 1 and 2) or not (Experiments 3 and 4). The results uniformly demonstrated that fast eye movements were completely stimulus driven, whereas slower eye movements were goal driven. These results are in line with neither a bottom-up account nor a top-down notion of visual selection. Instead, they indicate that visual selection is the outcome of 2 independent processes, one stimulus driven and the other goal driven, operating in different time windows. |
Lars Michels; Markus Lappe Contrast dependency of saccadic compression and suppression Journal Article In: Vision Research, vol. 44, no. 20, pp. 2327–2336, 2004. @article{Michels2004, In the occurrence of a saccadic eye movement vision becomes suppressed. Supra-threshold visual stimuli that are briefly presented at that time become perceptually compressed towards the saccade target (saccadic compression) and shifted in saccade direction (saccadic shift). We show that the strength of saccadic compression, like the strength of saccadic suppression, varies with stimulus contrast. Low contrast stimuli lead to stronger compression than high contrast stimuli. The similarity of contrast dependence and time course suggests that saccadic compression is related to saccadic suppression. Because the saccadic shift did not depend on contrast we suggest that shift and compression are different effects. |
Nadia Alahyane; Ansgar Koene; Denis Pélisson Transfer of adaptation from visually guided saccades to averaging saccades elicited by double visual targets Journal Article In: European Journal of Neuroscience, vol. 20, no. 3, pp. 827–836, 2004. @article{Alahyane2004a, The adaptive mechanisms that control the amplitude of visually guided saccades (VGS) are only partially elucidated. In this study, we investigated, in six human subjects, the transfer of VGS adaptation to averaging saccades elicited by the simultaneous presentation of two visual targets. The generation of averaging saccades requires the transformation of two representations encoding the desired eye displacement toward each of the two targets into a single representation encoding the averaging saccade (averaging programming site). We aimed to evaluate whether VGS adaptation acts upstream (hypothesis 1) or at/below (hypothesis 2) the level of averaging saccades programming. Using the double-step target paradigm, we simultaneously induced a backward adaptation of 17.5 degrees horizontal VGS and a forward adaptation of 17.5 degrees oblique VGS performed along the +/- 40 degrees directions relative to the azimuth. We measured the effects of this dual adaptation protocol on averaging saccades triggered by two simultaneous targets located at 17.5 degrees along the +/- 40 degrees directions. To increase the yield of averaging saccades, we instructed the subjects to move their eyes as fast as possible to an intermediate position between the two targets. We found that the amplitude of averaging saccades was smaller after VGS adaptation than before and differed significantly from that predicted by hypothesis 1, but not by hypothesis 2, with an adaptation transfer of 50%. These findings indicate that VGS adaptation largely occurs at/below the averaging saccade programming site. Based on current knowledge of the neural substrate of averaging saccades, we suggest that VGS adaptation mainly acts at the level of the superior colliculus or downstream. |
Nadia Alahyane; Denis Pélisson Eye position specificity of saccadic adaptation Journal Article In: Investigative Ophthalmology & Visual Science, vol. 45, no. 1, pp. 123–130, 2004. @article{Alahyane2004, PURPOSE: The accuracy of saccadic eye movements is maintained throughout life by adaptive mechanisms. With the double-step target paradigm, eight human subjects were investigated to determine whether saccadic adaptation depends only on the eye-displacement vector, or also on eye position as a context cue when two saccades of identical vector are adapted simultaneously. METHODS: First, bidirectional adaptations (BDAs) of horizontal saccades of the same vector were induced in a single training phase. Each direction of adaptation in BDAs (backward and forward) was linked to one vertical eye position (e.g., forward adaptation performed with the eyes directed 12.5 degrees upward and backward adaptation with the eyes 25 degrees downward) and alternated from trial to trial. Second, unidirectional adaptations (UDAs) were tested in two control conditions in which training trials of a single direction (backward or forward) were presented at both 12.5 degrees and -25 degrees eye elevations. RESULTS: Opposite changes in saccade amplitude could develop simultaneously in BDA, indicating that saccadic adaptation depends on orbital eye position. Comparing these data with the control conditions further indicated that eye position specificity was complete for backward, but not for forward, adaptation. CONCLUSIONS: The results indicate that saccadic adaptation mechanisms use vectorial eye displacement signals, but can also take eye position signals into account as a contextual cue when the training involves conflicting saccade amplitude changes. |
Jay Pratt; Jiye Shen; Jos J. Adam The planning and execution of sequential eye movements: Saccades do not show the one target advantage Journal Article In: Human Movement Science, vol. 22, no. 6, pp. 679–688, 2004. @article{Pratt2004, The present experiment examined the one-target advantage (OTA) with regard to saccadic eye movements. The OTA, previously found with manual pointing responses, refers to the finding that movements are executed faster when the limb is allowed to stop on the target compared to the situation where it has to proceed and hit a second target. Using an adapted limb movement OTA task, saccades of 5° and 15° were made to (a) a single target (one-target), (b) one target and immediately to another target without a change in direction (two-target-extension), and (c) one target and immediately back to the start location (two-target-reversal). Unlike manual movements, the movement times for the initial saccade in the two-target-extension condition were not prolonged compared to either of the other two conditions. Moreover, this pattern of results was found for both the shorter and longer amplitude saccades. The results indicate that the OTA does not occur in the oculomotor system and therefore is not a general motor control phenomenon. |
Eyal M. Reingold; Dave M. Stampe Saccadic inhibition in reading Journal Article In: Journal of Experimental Psychology: Human Perception and Performance, vol. 30, no. 1, pp. 194–211, 2004. @article{Reingold2004, In 5 experiments, participants read text that was briefly replaced by a transient image for 33 ms at random intervals. A decrease in saccadic frequency, referred to as saccadic inhibition, occurred as early as 60-70 ms following the onset of abrupt changes in visual input. It was demonstrated that the saccadic inhibition was influenced by the saliency of the visual event (Experiment 3) and was not produced in response to abrupt but irrelevant auditory stimuli (Experiment 1). Display changes restricted to an area either inside or outside the perceptual span required for normal reading produced strong saccadic inhibition (Experiment 2). Finally, Experiments 4 and 5 demonstrated higher level cognitive or attentional modulation of the saccadic inhibition effect. |
Millard Reschke; Jeffrey T. Somers; R. John Leigh; Jody M. Krnavek; Ludmila Kornilova; Inessa Kozlovskaya; Jacob J. Bloomberg; William H. Paloski Sensorimotor recovery following spaceflight may be due to frequent square-wave saccadic intrusions Journal Article In: Aviation Space and Environmental Medicine, vol. 75, no. 8, pp. 700–704, 2004. @article{Reschke2004, Square-wave jerks (SWJs) are small, involuntary saccades that disrupt steady fixation. We report the case of an astronaut (approximately 140 d on orbit) who showed frequent SWJs, especially postflight, but who showed no impairment of vision or decrement of postflight performance. These data support the view that SWJs do not impair vision because they are paired movements, consisting of a small saccade away from the fixation position followed, within 200 ms, by a corrective saccade that brings the eye back on target. Since many returning astronauts show a decrement of dynamic visual function during postflight locomotion, it seems possible that frequent SWJs improved this astronaut's visual function by providing postsaccadic enhancement of visual fixation, which aided postflight performance. Certainly, frequent SWJs did not impair performance in this astronaut, who had no other neurological disorder. |
D. C. Rijkaart; Josef N. Geest; Willem P. A. Kelders; Chris I. De Zeeuw; Maarten A. Frens Short-term adaptation of the cervico-ocular reflex Journal Article In: Experimental Brain Research, vol. 156, no. 1, pp. 124–128, 2004. @article{Rijkaart2004, The cervico-ocular reflex (COR) works in conjunction with the vestibulo-ocular reflex (VOR) and the optokinetic reflex (OKR) in order to prevent visual slip over the retina during head movement. The COR induces eye movements in response to proprioceptive signals from the neck. We investigated whether the COR gain can be adapted by inducing a mismatch between vision and neck proprioception, in analogy to VOR adaptation. Thirteen healthy subjects were rotated in the dark in a trunk-to-head manner (the head fixed in space while the body passively rotated sinusoidally with a peak velocity of 1.25 degrees /s). Eye movements were recorded with infrared video-oculography under various adaptive conditions. Analysis showed a small but significant reduction in COR gain in the suppression conditions. This means that the cervico-ocular reflex can be modified after only 10 min of concurrent visual and cervical stimulation. |
Martin Rolfs; Ralf Engbert Microsaccade orientation supports attentional enhancement opposite a peripheral cue Journal Article In: Psychological Science, vol. 15, no. 10, pp. 705–707, 2004. @article{Rolfs2004, Tse, Sheinberg, and Logothetis (2003) exploited a change-blindness paradigm to map the redistribution of spatial attention in response to a peripherally flashed cue. The probability of change detection at a given location was used as a measure of attention allocation. Using this measure, a ‘‘hot spot'' of attention (i.e., close to perfect change detection) was found along the cue-fixation axis. This hot spot ex- tended in the hemifield opposite the cued location. Here we show that an analysis of fixational eye movements in a spatial-cuing paradigm supports this important finding. During fixation of a stationary target, micromovements of the eyes occur involuntarily. These fixational eye movements are classified as drift, tremor, and microsaccades. Microsaccades are the fastest of these three components, with amplitudes typically smaller than 11 of visual angle. Using a classical spatial-cuing paradigm (Posner, 1980) with central arrow cues indicating the most probable location of a later target, we recently demonstrated that microsaccades are modulated by visual attention (Engbert & Kliegl, 2003b). About 300 ms after cue presentation, the angular distribution of microsaccades was oriented toward the expected target location. In the study reported here, we used pilot data from a larger ex- perimental study to examine the impact of peripheral cues on the rate and orientation of microsaccades. Using peripheral flashes—similar to the uninformative stimuli employed by Tse et al. (2003)—we found an orientation shift of microsaccades in the direction opposite the cued location. Taken together with our results on the coupling of visuo- spatial attention and microsaccades (Engbert & Kliegl, 2003b), these results are in agreement with the attentional enhancement opposite the cued location observed by Tse et al. (2003). |
Gerben Rotman; Eli Brenner; Jeroen B. J. Smeets Mislocalization of targets flashed during smooth pursuit depends on the change in gaze direction after the flash Journal Article In: Journal of Vision, vol. 4, pp. 564–574, 2004. @article{Rotman2004, Subjects mislocalize the position of a target that is flashed while they are making pursuit eye movements. This mislocalization is in the direction of pursuit. However, it is not clear whether it is the movement of the eyes or the movement of the pursuit target that matters. Neither is it clear whether it is the movement after the flash or the movement before the flash that matters. To resolve these issues, we asked subjects to pursue a disk that regularly changed its movement direction. Each change was followed by a change in the direction of gaze movement. Subjects were asked to tap targets that were flashed close to the moment at which the pursuit disk changed direction. We measured the movements of the eyes, head, and index finger. Subjects did not make saccades to the position they tapped but kept pursuing the disk. We compared the direction of the mislocalization with the changes in gaze and in target position during different intervals relative to the flash. We found that the mislocalization is related to the change in gaze after the flash. |
Sonja Stork; Jochen Müsseler Perceived localizations and eye movements with action-generated and computer-generated vanishing points of moving stimuli Journal Article In: Visual Cognition, vol. 11, no. 2-3, pp. 299–314, 2004. @article{Stork2004, When observers localize the vanishing point of a moving target, localizations are reliably displaced beyond the final position, in the direction the stimulus was travelling just prior to its offset. We examined modulations of this phenomenon through eye movements and action control over the vanishing point. In Experiment 1 with pursuit eye movements, localization errors were in movement direction, but less pronounced when the vanishing point was self-determined by a key press of the observer. In contrast, in Experiment 2 with fixation instruction, localization errors were opposite movement direction and independent from action control. This pattern of results points at the role of eye movements, which were gathered in Experiment 3. That experiment showed that the eyes lagged behind the target at the point in time, when it vanished from the screen, but that the eyes continued to drift on the targets' virtual trajectory. It is suggested that the perceived target position resulted from the spatial lag of the eyes and of the persisting retinal image during the drift. |
Hans Colonius; Adele Diederich Multisensory interaction in saccadic reaction time: A time-window-of-integration model Journal Article In: Journal of Cognitive Neuroscience, vol. 16, no. 6, pp. 1000–1009, 2004. @article{Colonius2004, Saccadic reaction time to visual targets tends to be faster when stimuli from another modality (in particular, audition and touch) are presented in close temporal or spatial proximity even when subjects are instructed to ignore the accessory input (focused attention task). Multisensory interaction effects measured in neural structures involved in saccade generation (in particular, the superior colliculus) have demonstrated a similar spatio-temporal dependence. Neural network models of multisensory spatial integration have been shown to generate convergence of the visual, auditory, and tactile reference frames and the sensorimotor coordinate transformations necessary for coordinated head and eye movements. However, because these models do not capture the temporal coincidences critical for multisensory integration to occur, they cannot easily predict multisensory effects observed in behavioral data such as saccadic reaction times. This article proposes a quantitative stochastic framework, the time-window-of-integration model, to account for the temporal rules of multisensory integration. Saccadic responses collected from a visual–tactile focused attention task are shown to be consistent with the time-window-of-integration model predictions. |
Ralf Engbert; Reinhold Kliegl Microsaccades keep the eyes' balance during fixation Journal Article In: Psychological Science, vol. 15, no. 6, pp. 431–436, 2004. @article{Engbert2004, During fixation of a stationary target, small involuntary eye movements exhibit an erratic trajectory—a random walk. Two types of these fixational eye movements are drift and microsaccades (small-amplitude saccades). We investigated fixational eye movements and binocular coordination using a statistical analysis that had previously been applied to human posture control. This random-walk analysis uncovered two different time scales in fixational eye movements and identified specific functions for microsaccades. On a short time scale, microsaccades enhanced perception by increasing fixation errors. On a long time scale, microsaccades reduced fixation errors and binocular disparity (relative to pure drift movements). Thus, our findings clarify the role of oculomotor processes during fixation. |
Jillian H. Fecteau; Crystal Au; Irene T. Armstrong; Douglas P. Munoz Sensory biases produce alternation advantage found in sequential saccadic eye movement tasks Journal Article In: Experimental Brain Research, vol. 159, no. 1, pp. 84–91, 2004. @article{Fecteau2004, In two-choice reaction time tasks, participants respond faster when the correct decision switches across consecutive trials. This alternation advantage has been interpreted as the guessing strategies of participants. Because the participants expect that the correct decision will switch across consecutive trials, they respond faster when this expectation is confirmed and they respond more slowly when it is disconfirmed. In this study, we evaluated the veracity of this expectancy interpretation. After replicating a long-lasting alternation advantage in saccadic reaction times (Experiment 1), we show that reducing the participants' ability to guess with a challenging mental rotation task does not change the alternation advantage, which suggests that expectancy is not responsible for the effect (Experiment 2). Next, we used prosaccade and antisaccade responses to dissociate between the sensory and motor contributions of the alternation advantage (Experiment 3) and we found that the alternation advantage originates from sensory processing. The implications of these findings are discussed with regard to guessing strategies, sensory processing, and how these findings may relate to inhibition of return. |
Giovanni Galfano; Elena Betta; Massimo Turatto Inhibition of return in microsaccades Journal Article In: Experimental Brain Research, vol. 159, no. 3, pp. 400–404, 2004. @article{Galfano2004, Inhibition of return (IOR) is the term used to describe the phenomenon whereby stimuli appearing at recently attended locations are reacted to less efficiently than stimuli appearing at locations that have not yet been attended. In the present study, we employed a typical IOR paradigm with peripheral uninformative cues while participants maintained their eyes at fixation. Eye position was monitored at a high sampling rate (500 Hz) in order to detect miniature eye movements called microsaccades, which have been shown to be crucial for avoiding disappearance of visual image. However, recent studies have demonstrated a close relationship between covert endogenous attentional shifts and the direction of microsaccades. Here, we demonstrate that the direction of microsaccades can be biased away from the peripheral location occupied by a salient, although task-irrelevant, visual signal. Because microsaccades are known not to be under conscious control, our results suggest strong links between IOR and unconscious oculomotor programming. |
2003 |
Ralf Engbert; Reinhold Kliegl Microsaccades uncover the orientation of covert attention Journal Article In: Vision Research, vol. 43, no. 9, pp. 1035–1045, 2003. @article{Engbert2003, Fixational eye movements are subdivided into tremor, drift, and microsaccades. All three types of miniature eye movements generate small random displacements of the retinal image when viewing a stationary scene. Here we investigate the modulation of microsaccades by shifts of covert attention in a classical spatial cueing paradigm. First, we replicate the suppression of microsaccades with a minimum rate about 150 ms after cue onset. Second, as a new finding we observe microsaccadic enhancement with a maximum rate about 350 ms after presentation of the cue. Third, we find a modulation of the orientation towards the cue direction. These multiple influences of visual attention on microsaccades accentuate their role for visual information processing. Furthermore, our results suggest that microsaccades can be used to map the orientation of visual attention in psychophysical experiments. |
Valérie Gaveau; Olivier Martin; Claude Prablanc; Denis Pélisson; Christian Urquizar; Michel Desmurget On-line modification of saccadic eye movements by retinal signals Journal Article In: Neuroreport, vol. 14, no. 6, pp. 875–878, 2003. @article{Gaveau2003, A saccade is a rapid shift of the position of the eyes (<100ms). Saccades are generally considered too quick to be inpoundsuenced by retinal signals.To address this idea, we displaced the visual target of a rightward horizontal saccade at eye movement onset (when there is suppression of conscious perception).To prevent adaptive and learning e¡ects to occur, jump saccadeswere always followed by a random series of 10 no-jump saccades. Results indicated that the target jump influenced significantly the amplitude and the peak velocity of the ongoing saccade (opposite e¡ects were found for rightward and leftward jumps). Changes in saccade kinematics occurred as early as 50ms after the target jump. These results show that retinal information is processed quickly during eye movements, presumably through sub-cortical pathways. |
Casimir J. H. Ludwig; Iain D. Gilchrist Target similarity affects saccade curvature away from irrelevant onsets Journal Article In: Experimental Brain Research, vol. 152, no. 1, pp. 60–69, 2003. @article{Ludwig2003, Saccade curvature away from visual distractors is a measure of the salience of these distractors for the oculomotor system. Three experiments are reported in which the integration of luminance onset signals and target similarity signals is examined, using a saccade curvature paradigm. Observers made saccades to a no-onset colour target in one of two positions on the vertical meridian. On most trials, an abrupt onset distractor that was either similar or dissimilar to the target appeared left or right on the horizontal midline. Saccades curved away from the irrelevant onsets; however, the amount of curvature was modulated by target similarity only when the onset appeared before the target (experiment 2) or when saccade initiation was delayed (experiment 3). These results suggest that the initial response to the onset is stimulus-driven and mediated by its transient component. Over time, the response is integrated with and augmented by top-down inputs. Visual and non-visual signals converge onto a common motor map to determine an item's salience. |
Benjamin T. Backus; Daniel Matza-Brown The contribution of vergence change to the measurement of relative disparity Journal Article In: Journal of Vision, vol. 3, no. 11, pp. 727–750, 2003. @article{Backus2003, The relative disparity between two objects in a scene can in principle be measured directly from the retinal images, without knowledge of eye position. But relative disparity increment thresholds are lowest when the relative disparity is small and the objects are not widely separated in the visual field: thus, some relative disparities are easier for the visual system to measure than others. We consider, after others, a second method by which the visual system could measure relative disparity, based on change in vergence ("delta vergence" or DV). The DV mechanism could be more reliable than the retinal mechanism when visual targets are widely separated in visual direction or depth. We used a cue-conflict paradigm to measure the extent to which perceived depth depends on DV. As target separation increased, so did reliance on DV. As intertarget disparity increased, reliance on DV increased for one observer but not for two others. |
Dominic J. Mort; Richard J. Perry; Sabira K. Mannan; Timothy L. Hodgson; Elaine Anderson; Rebecca Quest; Donald McRobbie; Alan McBride; Masud Husain; Christopher Kennard Differential cortical activation during voluntary and reflexive saccades in man Journal Article In: NeuroImage, vol. 18, no. 2, pp. 231–246, 2003. @article{Mort2003, A saccade involves both a step in eye position and an obligatory shift in spatial attention. The traditional division of saccades into two types, the "reflexive" saccade made in response to an exogenous stimulus change in the visual periphery and the "voluntary" saccade based on an endogenous judgement to move gaze, is supported by lines of evidence which include the longer onset latency of the latter and the differential effects of lesions in humans and primates on each. It has been supposed that differences between the two types of saccade derive from differences in how the spatial attention shifts involved in each are processed. However, while functional imaging studies have affirmed the close link between saccades and attentional shifts by showing they activate overlapping cortical networks, attempts to contrast exogenous with endogenous ("covert") attentional shifts directly have not revealed separate patterns of cortical activation. We took the "overt" approach, contrasting whole reflexive and voluntary saccades using event-related fMRI. This demonstrated that, relative to reflexive saccades, voluntary saccades produced greater activation within the frontal eye fields and the saccade-related area of the intraparietal sulci. The reverse contrast showed reflexive saccades to be associated with relative activation of the angular gyrus of the inferior parietal lobule, strongest in the right hemisphere. The frequent involvement of the right inferior parietal lobule in lesions causing hemispatial neglect has long implicated this parietal region in an important, though as yet uncertain, role in the awareness and exploration of space. This is the first study to demonstrate preferential activation of an area in its posterior part, the right angular gyrus, during production of exogenously triggered rather than endogenously generated saccades, a finding which we propose is consistent with an important role for the angular gyrus in exogenous saccadic orienting. |
Nadia Alahyane; Denis Pélisson Adaptation of saccadic eye movements: Transfer and specificity Journal Article In: Annals of the New York Academy of Sciences, vol. 1004, no. 1, pp. 69–77, 2003. @article{Alahyane2003, The present study was designed to test whether the adaptation of saccadic eye movements depends only on the eye displacement vector of the trained saccade or also on eye position information. Using the double-step target paradigm in eight human subjects, we first induced in a single session two "opposite directions adaptations" (ODA) of horizontal saccades of the same vector. Each ODA (backward or forward) was linked to one vertical eye position (12.5 degrees up or 25 degrees down) and alternated from trial to trial. The results showed that opposite changes of saccade amplitude can develop simultaneously, indicating that saccadic adaptation depends on orbital eye position. This finding has important functional implications because in everyday life our eyes saccade from constantly changing orbital positions. A comparison of these data to two control conditions in which training trials of a single type (backward or forward) were presented at both 12.5 degrees and -25 degrees eye elevations further indicated that eye position specificity is complete for backward, but not for forward, adaptation. Finally, the control conditions also indicated that the adaptation of a single saccade fully transferred to untrained saccades of the same vector, but initiated from different vertical eye positions. In conclusion, our study indicates that saccadic adaptation mechanisms use vectorial eye displacement signals, but can also take eye position signals into account as a contextual cue when the training involves conflicting saccade amplitude changes |
Richard Amlôt; Robin Walker; Jon Driver; Charles Spence Multimodal visual–somatosensory integration in saccade generation Journal Article In: Neuropsychologia, vol. 41, no. 1, pp. 1–15, 2003. @article{Amlot2003, Neurophysiological studies have demonstrated multisensory interaction effects in the neural structures involved in saccade generation when visual, auditory or somatosensory stimuli are presented bimodally. Visual–auditory interaction effects have been demonstrated in numerous behavioural studies of saccades but little is known about interaction effects involving somatosensory stimuli. The present study examined visual–somatosensory interaction effects on saccade generation using a multisensory paradigm, whereby task-irrelevant distractors appeared spatially-coincident with, or remote from the designated saccade target. Somatosensory distractors reduced the latency of saccades when presented before the visual target and the greatest facilitation effectwas observed with spatially-coincident stimuli.Visual distractors spatially-coincident with a somatosensory target reduced latency (and increased peak velocity) when presented before and after the target.Visual distractors contralateral to somatosensory targets increased saccade latency and produced high error rates of saccades made to the distractor. The high error rates and latencymodulation with visual distractors is consistent with a bias for visual stimuli in the saccadic system. In the visual target condition, saccade latency was modulated by a somatosensory distractor that was entirely task-irrelevant and this effect was always greatest with spatially-coincident distractors. The multisensory distractor effects are discussed in terms of saccades being programmed to the non-target modality, the early triggering of a non-spatial saccade ‘when' signal, and multisensory neuronal enhancement effects. |
Ryota Kanai; Josef N. Geest; Maarten A. Frens Inhibition of saccade initiation by preceding smooth pursuit Journal Article In: Experimental Brain Research, vol. 148, no. 3, pp. 300–307, 2003. @article{Kanai2003, In this study, we investigated the influence of smooth-pursuit eye movements on saccade initiation in response to a sudden jump of a continuously moving target. We replicated the finding by Tanaka et al. (1998) that saccadic eye movements in the direction opposite to preceding pursuit have longer latencies than those in the same direction. We confirmed that this asymmetry is indeed due to an inhibitory effect of smooth pursuit on saccade initiation in the opposite direction rather than facilitation of saccade initiation in the same direction. The inhibitory effect decreased strongly when subjects knew the jump direction in advance. This supports the notion that the prolonged latencies of backward saccades are not due to orbital mechanics or low-level motor processing. Furthermore, we found that the range of saccade direc- tions inhibited by a pursuit movement is broad, covering all directions that did not have the same horizontal component as the pursuit direction. This is in contrast with the predictions of “Inhibition of Saccade Return” (ISR, Hooge and Frens 2000), which is restricted to a smaller confined area. |
Adele Diederich; Hans Colonius; Daniela Bockhorst; Sandra Tabeling Visual-tactile spatial interaction in saccade generation Journal Article In: Experimental Brain Research, vol. 148, no. 3, pp. 328–337, 2003. @article{Diederich2003, Saccadic reaction times to visual targets tend to be faster when non-visual stimuli are presented in close temporal or spatial proximity even if subjects are instructed to ignore the accessory input. The effect tends to decrease with increasing spatial distance between the stimuli. Multisensory interaction effects measured in neural structures involved in saccade generation have demonstrated a similar spatial dependence. The present study investigated visual-tactile interaction effects on saccadic reaction time using a focused attention paradigm. Compared to unimodal visual targets saccadic reaction time to bimodal stimuli was reduced by up to 30 ms. The effect was larger for ipsi- than for contralateral presentations, and it increased with the eccentricity of the visual target. The results are consistent with attributing part of the facilitation to a multisensory effect of bimodal neurons with overlapping visual and tactile receptive field structures in the deep layers of the superior colliculus. |
Jeroen B. J. Smeets; Ignace T. C. Hooge Nature of variability in saccades Journal Article In: Journal of Neurophysiology, vol. 90, no. 1, pp. 12–20, 2003. @article{Smeets2003, We studied the variability in saccades by comparing the peak velocities of saccades with the same target amplitude made with different actual amplitudes. We tested three hypotheses: the pulse-height noise hypothesis (peak velocity and amplitude vary proportionally), the localization noise hypothesis (variability in amplitude and peak velocity lie along the main sequence), and the independent noise hypothesis (variability in amplitude and peak velocity are independent). We measured eye orientation in two experiments by a scleral coil and a video system. Surprisingly, the main source of variability of saccades depended on the measurement system used. A combination of localization noise and independent noise best describes the data obtained by the video system. The independent noise (e.g., measurement inaccuracy) was the main source of variability. For the scleral coils, the variability was considerably larger than for the less accurate video system. The pulse-height noise hypothesis best describes this additional variability. Therefore we conclude that pulse-height noise is the main source of variability in saccades measured with scleral coils. We discuss the influence of scleral coils on saccade generation and suggest that a change in motor strategy due to the discomfort of wearing the coils might be the cause of the increased variability. |
2002 |
Sonja Stork; Sebastiaan F. W. Neggers; Jochen Müsseler Intentionally-evoked modulations of smooth pursuit eye movements Journal Article In: Human Movement Science, vol. 21, no. 3, pp. 335–348, 2002. @article{Stork2002, When observers pursue a moving target with their eyes, they use predictions of future target positions in order to keep the target within the fovea. It was suggested that these predictions of smooth pursuit (SP) eye movements are computed only from the visual feedback of the target characteristics. As a consequence, if the target vanishes unexpectedly, the eye movements do not stop immediately, but they overshoot the vanishing point. We compared the spatial and temporal features of such predictive eye movements in a task with or without intentional control over the target vanishing point. If the observers stopped the target with a button press, the overshoot of the eyes was reduced compared to a condition where the offset was computer generated. Accordingly, the eyes started to decelerate well before the target offset and lagged further behind the target when it disappeared. The involvement of intentionally-generated expectancies in eye movement control was also obvious in the spatial trajectories of the eyes, which showed a clear flexion in anticipation of the circular motion path we used. These findings are discussed together with neurophysiological mechanisms underlying the SP eye movements. |
Eyal M. Reingold; Dave M. Stampe Saccadic inhibition in voluntary and reflexive saccades Journal Article In: Journal of Cognitive Neuroscience, vol. 14, no. 3, pp. 371–388, 2002. @article{Reingold2002b, The present study investigated saccadic inhibition in both voluntary and stimulus-elicited saccades. Two experiments examined saccadic inhibition caused by an irrelevant flash occurring subsequent to target onset. In each trial, participants were required to perform a single saccade following the presentation of a black target on a gray background, 4 degrees to the left or to the right of screen center. In some trials (flash trials), after a variable delay, a 33-msec flash was displayed at the top and bottom third of the monitor (these regions turned white). In all experimental conditions, histograms of flash-to-saccade latencies documented a decrease in saccadic frequency, forming a dip, time-locked to the flash and occurring as early as 60-70 msec following its onset. The fast latency of this effect strongly suggests a low-level, reflex-like, oculomotor effect, which was referred to as saccadic inhibition. A novel procedure was developed to allow comparisons of saccadic inhibition even across conditions, which in the absence of a flash (no-flash trials) produce dissimilar saccadic reaction times (SRTs) distributions. Experiment 1 examined the effects of the fixation stimulus on saccadic inhibition by contrasting three conditions: a gap condition (fixation stimulus disappeared 200 msec prior to target onset), a step condition (offset of the fixation stimulus was simultaneous with target onset), and an overlap condition (the fixation stimulus remained on for the duration of the trial). The overlap condition produced substantially stronger saccadic inhibition, relative to the gap and the step conditions. Experiment 2 contrasted the saccadic inhibition effect obtained for prosaccades (saccades aimed at the target) with the effect obtained for antisaccades (i.e., saccades aimed away from the same target). The onset of saccadic inhibition was earlier, and its magnitude was stronger, for antisaccades, relative to prosaccades. The plausibility that the superior colliculus is the neurophysiological locus of the saccadic inhibition effect was explored. |
F. Møller; M. L. Laursen; J. Tygesen; A. K. Sjølie Binocular quantification and characterization of microsaccades Journal Article In: Graefe's Archive for Clinical and Experimental Ophthalmology, vol. 240, no. 9, pp. 765–770, 2002. @article{Moeller2002, BACKGROUND: The significance of microsaccades in the visual process has been discussed for more than 50 years. However, only a few studies have measured microsaccades binocularly, and detailed quantification and characterization of these small movements are needed in order to further understand their nature. METHOD: The amplitude, velocity, acceleration and direction of microsaccades were quantified binocularly in 10 normal test persons during a 40-s fixation task, using an infrared recording technique. RESULTS: All microsaccades for all test persons were performed simultaneously and individually with an almost identical amplitude in the right and left eye (a range of 0.003-0.042 deg between right and left eye mean values). The mean microsaccadic amplitude for the test persons was within a range of 0.223-1.079 deg. The directional difference between simultaneously-performed right and left eye microsaccades was less than 22.5 deg for 84.8% of the saccades, indicating that the majority of microsaccades are conjugated. Three different fixation patterns were identified and characterized: (1) a classic interplay between easily identified drifts and medium-sized microsaccades (mean amplitude range 0.328-0.413 deg); (2) long intersaccadic intervals (4-5 s) with almost absent drifts, followed by three or four large microsaccades (mean amplitude range 0.755-1.079 deg); and (3) low-amplitude drift movements interrupted by low-amplitude microsaccades (mean amplitude range 0.231-0.265 deg). CONCLUSION: Microsaccades are involuntary, predominantly conjugated, simultaneously performed, and of almost identical amplitude in the right and left eye, suggesting a central control mechanism for microsaccades at subcortical level. |
Danny Gagnon; Gillian A. O'Driscoll; Michael Petrides; G. B. Pike The effect of spatial and temporal information on saccades and neural activity in oculomotor structures Journal Article In: Brain, vol. 125, no. 1, pp. 123–139, 2002. @article{Gagnon2002, It has been argued that saccade generation is supported by two systems, a'where' system that decides the direction and extent of an impending saccade, and a 'when' system that is involved in the timing of the release of fixation. We evaluated the contributions of these systems to saccade latencies, and used functional MRI to identify the neural substrates of these systems. We found that advance knowledge of the direction and the timing of an impending target movement had both overlapping and discrete effects on saccade latencies and on neural activation. Knowledge of either factor decreased regular saccade latencies. However, knowledge of target direction increased the number of predictive and express saccades while knowledge of target timing did not. The brain activation data showed that advance knowledge of the direction or the timing of the target movement activated primarily overlapping structures. The precentral gyrus, in the region of the frontal eye fields, was more active in conditions in which some aspect of the target movement was predictable than in saccade control and fixation conditions. In the basal ganglia, activation discriminated between advance knowledge of target timing and target direction. The lenticular nuclei were more active when only target timing was known in advance, while the caudate was more active when only target direction was known in advance. These data suggest that the neural structures supporting the 'where' and 'when' systems are highly overlapping, although there is some dissociation sub-cortically. Knowledge of target timing and target direction converge in precentral gyrus, a region where there is strong evidence of context-dependent modulation of neural activity. |
Richard Godijn; Jan Theeuwes Programming of exogenous and endogenous saccades: Evidence for a competitive integration model Journal Article In: Journal of Experimental Psychology: Human Perception and Performance, vol. 28, pp. 1039–1054, 2002. @article{Godijn2002, Participants were required to make a saccade to a uniquely colored target while ignoring the presentation of an onset distractor. The results provide evidence for a competitive integration model of saccade programming that assumes endogenous and exogenous saccades are programmed in a common saccade map. The model incorporates a lateral interaction structure in which saccade-related activation at a specific location spreads to neighboring locations but inhibits distant locations. In addition, there is top-down, location-specific inhibition of locations to which the saccade should not go. The time course of exogenous and endogenous activation in the saccade map can explain a variety of eye movement data, including endpoints, latencies, and trajectories of saccades and the well-known global effect |
Martin Greschner; Markus Bongard; Pal Rujan; Josef Ammermüller Retinal ganglion cell synchronization by fixational eye movements improves feature estimation Journal Article In: Nature Neuroscience, vol. 5, no. 4, pp. 341–347, 2002. @article{Greschner2002, Image movements relative to the retina are essential for the visual perception of stationary objects during fixation. Here we have measured fixational eye and head movements of the turtle, and determined their effects on the activity of retinal ganglion cells by simulating the movements on the isolated retina. We show that ganglion cells respond mainly to components of periodic eye movement that have amplitudes of roughly the diameter of a photoreceptor. Drift or small head movements have little effect. Driven cells that are located along contrast borders are synchronized, which reliably signals a preceding movement. In an artificial neural network, the estimation of spatial frequencies for various square wave gratings improves when timelocked to this synchronization. This could potentially improve stimulus feature estimation by the brain. |
Casimir J. H. Ludwig; Iain D. Gilchrist Measuring saccade curvature: A curve-fitting approach Journal Article In: Behavior Research Methods, Instruments & Computers, vol. 34, no. 4, pp. 618–624, 2002. @article{Ludwig2002, Saccade curvature is becoming a popular measure for detecting the presence of competing saccadic motor programs. Several different methods of quantifying saccade curvature have been employed. In the present study, we compared these metrics with each other and with novel measures based on curve fitting. Initial deviation metrics were only moderately associated with the more widely used metric of maximum curvature. The latter was strongly related to a recently developed area-based measure and to the novel methods based on second- and third-order polynomial fits. The curve-fitting methods showed that although most saccades curved in only one direction, there was a population of trajectories with both a maximum and a minimum (i.e., double-curved saccades). We argue that a curvature metric based on a quadratic polynomial fit deals effectively with both types of trajectories and, because it is based on all the samples of a saccade, is less susceptible to sampling noise. |
M. C. Doyle; Robin Walker Multisensory interactions in saccade target selection: Curved saccade trajectories Journal Article In: Experimental Brain Research, vol. 142, no. 1, pp. 116–130, 2002. @article{Doyle2002, In a series of experiments, we examined the change in saccade trajectories observed when distractors are presented at non-target locations. The primary aim of the experiments was to examine multisensory interaction effects between the visual, auditory and somatosensory modalities in saccade generation. In each experiment observers made saccades to visual targets above and below fixation in the presence of visual, auditory or tactile stimuli to the left or right of fixation. In experiment 1 distractor location indicated which of two stimuli was the target for the saccade. Saccade trajectories showed strong leftward curvature following right-side distractors and showed rightward curvature following left-side distractors. The largest effects on trajectories were observed for visual distractors, but significant curvature was observed with auditory and somatosensory distractors. In experiment 2 saccades were made following the onset of a visual target (reflexive) or following presentation of an arrow at fixation (voluntary), and task-irrelevant crossmodal distractors were presented simultaneously with target onset. Both voluntary and reflexive saccades were found to curve away from task-irrelevant visual distractors, but auditory and somatosensory distractors did not modulate saccade trajectories. In experiment 3 task-irrelevant distractors preceded the onset of the target by 100 ms. Reflexive saccades were found to curve away from visual, auditory and somatosensory distractors, but voluntary saccades curved away from visual distractors only. The modulation of saccade trajectories by distractors from different modalities is interpreted in terms of inhibitory processes operating in neural structures involved in saccade generation. Our findings suggest that visual, auditory and somatosensory distractors can all modulate saccade trajectories. Such effects could be related to the inhibition of populations of neurons, in a common motor map, for the selection of a saccade target. |
Veerle Gysen; Peter De Graef; Karl Verfaillie Detection of intrasaccadic displacements and depth rotations of moving objects Journal Article In: Vision Research, vol. 42, no. 3, pp. 379–391, 2002. @article{Gysen2002, In a display with a stationary and a moving object, subjects saccaded towards one of the objects and had to detect intrasaccadic changes in position or orientation of either the saccade target or the saccade flanker. Compared to performance for stationary objects, displacement detection for translating objects was better and unaffected by saccadic status of the changed object. This pattern proved to be specific to position changes in translating objects and did not generalize to other types of motion (i.e., rotation) or to other types of intrasaccadic changes (i.e., orientation shifts). Superior transsaccadic coding of the position of a translating object was also observed in control experiments with only a single object present on each trial. Possible accounts in terms of selective attention to moving objects and perceptual relevance of object position are pitted against the data, suggesting qualitative differences in the transsaccadic representation of translating and stationary objects. |
Monika Harvey; Bettina Olk; Keith Muir; Iain D. Gilchrist Manual responses and saccades in chronic and recovered hemispatial neglect: A study using visual search Journal Article In: Neuropsychologia, vol. 40, no. 7, pp. 705–717, 2002. @article{Harvey2002, Hemispatial neglect affects both the ability to respond to targets on the contralesional side of space and to programme saccades to such targets. In the current study, we looked in detail at saccade programming and manual reaction times (RTs) in a range of visual search tasks, in which task difficulty was systematically increased by changing the nature of the distractors. In condition 1, the target was presented with no distractors. In the other conditions, displays contained three distractors that were changed across conditions to manipulate similarity to the target and so task difficulty. We tested two neglect patients, one chronic, one recovered along with two RCVA control patients and 12 age-matched controls. Both neglect patients studied could successfully execute saccades into the neglected field when the target was presented alone. However, a dissociation emerged between the two patients when the target was presented with distractor items. Patient ERs first saccade to target performance in the three search conditions revealed clear effects of distractor type. In contrast for the recovered patient AF, the left/right difference was present for all search displays and appeared to be constant regardless of distractor type. This differential pattern of behaviour may reflect the different underlying neural causes of the neglect in these patients. In the current study, the measurement of saccades allowed the task to be fractionated, and thus, reveal the action of multiple mechanisms controlling saccades in search. |
Hyung-Chul O. Li; Eli Brenner; Frans W. Cornelissen; Eun-Soo Kim Systematic distortion of perceived 2D shape during smooth pursuit eye movements Journal Article In: Vision Research, vol. 42, no. 23, pp. 2569–2575, 2002. @article{Li2002b, Even when the retinal image of a static scene is constantly shifting, as occurs when the viewer pursues a small moving object with his or her eyes, the scene is usually correctly perceived to be static. Following early suggestions by von Helmholtz, it is commonly believed that this spatial stability is achieved by combining retinal and extra-retinal signals. Here, we report a perceptually salient 2D shape distortion that can arise during pursuit. We provide evidence that the perceived 2D shape reflects retinal image contents alone, implying that the extra-retinal signal is ignored when judging 2D shape. © 2002 Elsevier Science Ltd. All rights reserved. |
2001 |
Sebastian Pannasch; Sascha M. Dornhoefer; Pieter J. A. Unema; Boris M. Velichkovsky The omnipresent prolongation of visual fixations: Saccades are inhibited by changes in situation and in subject's activity Journal Article In: Vision Research, vol. 41, no. 25-26, pp. 3345–3351, 2001. @article{Pannasch2001, Presenting a distractor prolongs not only saccadic reaction times in paced tasks but also fixation durations in unpaced tasks. To investigate whether the effect of a distractor is a pure optomotor reflex, we used both visual and auditory distractors in an unpaced picture-viewing paradigm. Results show a distractor effect for both modalities. Analysis of data from previous studies showed similar effects, even in amodal shifts of attention. These findings challenge the hypothesis that the effect is modality-specific and suggest that the distractor effect may be another expression of the orienting reflex. |
Risto Näsänen; Helena Ojanpää; Ilpo Kojo Effect of stimulus contrast on performance and eye movements in visual search Journal Article In: Vision Research, vol. 41, no. 14, pp. 1817–1824, 2001. @article{Naesaenen2001a, According to the visual span control hypothesis, eye movements are controlled in relation to the size of visual span. In reading, the decrease of contrast reduces visual span, saccade sizes, and reading speed. The purpose of the present study is to determine how stimulus contrast affects the speed of two-dimensional visual search and how changes in eye movements and visual span could explain changes in performance. The task of the observer was to search for, and identify, an uppercase letter from a rectangular array of characters in which the other items were numerals. Threshold search time, i.e. the duration of stimulus presentation required for search that is successful with a given probability, was determined by using a multiple-alternative staircase method. Eye movements were recorded simultaneously by using a video eye tracker. Four different set sizes (the sizes of stimulus array) (3 × 3-10 × 10), and five different contrasts (0.0186 - 0.412) were used. At all set sizes, threshold search time decreased with increasing contrast. Also the average number of fixations per search decreased with increasing contrast. At the smallest set size (3 × 3), only one fixation was needed except at the lowest contrast. Average fixation duration decreased and saccade amplitudes increased slightly with increasing contrast. The reduction of the number of fixations with increasing contrast suggests that visual span, i.e. the area from which information can be collected at one fixation, increases with increasing contrast. The reduction of the number of fixations together with reduced fixation duration result in reduced search times when contrast increases. |
Albert V. Berg; J. A. Beintema; Maarten A. Frens Heading and path percepts from visual flow and eye pursuit signals Journal Article In: Vision Research, vol. 41, no. 25-26, pp. 3467–3486, 2001. @article{Berg2001, The percept of self-motion through the environment is supported by visual motion signals and eye movement signals. The interaction between these signals by decoupling of the eye movement and the pattern of retinal motion during brief simulated ego-movement on straight or circular trajectories was studied. A new response method enabled subjects to report perceived destination and perceived curvature of their future path simultaneously. Various combinations of simulated gaze rotation in the retinal flow and eye pursuit were investigated. Simulated gaze rotation ranged from consistent and larger than, to opponent and larger than eye pursuit. It was found that the perceived destination shifts non-linearly with the mismatch between simulated gaze rotation and eye pursuit. The non-linearity is also revealed in the perceived tangent heading direction and perceived path curvature, although to different extent in different subjects. For the same retinal flow, eye pursuit that is consistent with the simulated gaze rotation reduces heading error and the perceived path straightens out. In contrast, perceived path and/or heading do not become more curved or more biased in the direction opposite to pursuit when the eye -in-head rotation is opposite to the simulated gaze rotation. These observations point to modulation of the effect of the extra-retinal pursuit signal by the visual evidence for eye rotation. In a second experiment, one presented to a stationary eye the sum of a component of simulated gaze rotation and radial flow. It was found that the bi-circular flow component, that characterizes the change in pattern of flow directions by the gaze rotation, induces a shift of perceived heading without appreciable perceived path curvature. Conversely, the complementary component of simulated gaze rotation (bi-radial flow) evokes a percept of motion on a curved path with a small tangent heading error. It was suggested that bi-circular and bi-radial flow components contribute primarily to percepts of heading and path curvature, respectively. |
Melanie Doyle; Robin Walker Curved saccade trajectories: Voluntary and reflexive saccades curve away from irrelevant distractors Journal Article In: Experimental Brain Research, vol. 139, no. 3, pp. 333–344, 2001. @article{Doyle2001, In this study we examined the impact of irrelevant distractors upon trajectories of reflexive and voluntary saccades. Observers made saccades to visual targets above and below fixation as directed by target appearance (reflexive) or by a central directional cue (voluntary) in the presence of an irrelevant distractor stimulus (a cross) whose appearance was simultaneous with target onset. We recorded saccade latency, amplitude and the magnitude of saccade curvature relative to the direct route from the start-to-end of the saccade. Previous studies of saccades curvature have used distractors to provide information about the saccade task and, as a result, have only examined trajectories of voluntary saccades. However, we have shown that both reflexive and voluntary saccades curved away from irrelevant distractors. The effect of irrelevant distractors indicates that observers do not need to attend to distractors in a voluntary fashion for distractors to modify saccade trajectories. Furthermore, it highlights an important parallel in curvature of saccades and reach trajectories, namely that both curve away from irrelevant distractors. The second important observation was that reflexive, as well as voluntary, saccades curved away from distractors. This suggests that curvature is not solely a consequence of voluntary control. These results have been considered within the context of inhibition-based theories of curvature derived from studies of saccade and manual reach trajectories. |
2000 |
Diane C. Gooding; Meghan D. Miller; Thomas R. Kwapil Smooth pursuit eye tracking and visual fixation in psychosis-prone individuals Journal Article In: Psychiatry Research, vol. 93, no. 1, pp. 41–54, 2000. @article{Gooding2000a, Subjects identified by Perceptual Aberration-Magical Ideation (Per-Mag) scores (n = 97), Social Anhedonia (SocAnh) scores (n = 45), and Physical Anhedonia (PhysAnh) scores (n = 31) as well as normal controls (n = 94), underwent psychophysiological and clinical assessment. This is the first published investigation of pursuit system functioning in three groups of questionnaire-identified at-risk individuals. Pursuit during a simple non- monitor tracking task was measured using root-mean-square error (RMSE) scores and pursuit gain scores. Fixation performance was measured in terms of number of saccades away from the central fixation point. The at-risk subjects were more likely to display aberrant smooth pursuit tracking than controls, though there were no significant differences between the at-risk subjects endorsing items relevant to positive-symptom schizotypy and those endorsing items pertaining to negative-symptom schizotypy. The groups did not differ significantly in their visual fixation performance. Participants were also evaluated for the presence of Axis I symptomatology and psychotic-like experiences. Neither the experimental subjects nor the control subjects displayed a significant association between ocular motor performance and psychotic-like experiences. These findings are consistent with prior evidence that pursuit tracking is a trait characteristic, independent of clinical status. |