During active vision, the eyes continually scan the visual environment using saccadic scanning movements. This target article presents an information processing model for the control of these movements, with some close parallels to established physiological processes in the oculomotor system. Two separate pathways are concerned with the spatial and the temporal programming of the movement. In the temporal pathway there is spatially distributed coding and the saccade target is selected from a “salience map.” Both pathways descend through a hierarchy of levels, the lower ones operating automatically. Visual onsets have automatic access to the eye control system via the lower levels. Various centres in each pathway are interconnected via reciprocal inhibition. The model accounts for a number of well-established phenomena in target-elicited saccades: the gap effect, express saccades, the remote distractor effect, and the global effect. High-level control of the pathways in tasks such as visual search and reading is discussed; it operates through spatial selection and search selection, which generally combine in an automated way. The model is examined in relation to data from patients with unilateral neglect.
In a series of experiments, we examined the increase in saccade latency that is observed consistently when distractor stimuli are presented simultaneously with the saccade target at various nontarget locations. In the first experiment, targets and distractors were presented on the horizontal axis. We found that saccade latency was increased when distractors appeared at fixation and in the contralateral nontarget hemifield (at eccentricities < or = 10 degrees). In contrast, latency was unaffected by distractors presented along the ipsilateral target axis, but amplitude was increased as saccades tended to land at intermediate locations between the two stimuli (global effect). The effect of presenting distractors at various two-dimensional locations in both the target and nontarget hemifields then was examined, and the maximum latency increase again was observed when distractors appeared at fixation. Distractors presented on any of the eight principal axes in either hemifield, other than on the horizontal target axis, also increased latency. The relationship between the effects of distractors on latency and amplitude was reciprocal. Within approximately 20 degrees of the target axis itself, distractors affected saccade amplitude but not latency. In contrast, distractors presented outside this "window" increased saccade latency without affecting amplitude. A systematic quantitative relationship was revealed between the increase in latency and the ratio between target and distractor eccentricities. The latency increase was largest with small values of the ratio and reached a peak with distractors at the fixation location. The finding that the increase observed for more eccentric distractor locations fitted the same function as that at fixation shows that inhibitory effects operate over large areas of the visual field. The increase in latency under distractor conditions is interpreted in light of recent neurophysiological findings of inhibitory processes operating in the rostral region of the superior colliculus. Our results suggest that these inhibitory processes are not restricted to the central foveal region alone but operate over wider regions of the visual field.
McSorley, Eugene, Patrick Haggard, and Robin Walker. Time course of oculomotor inhibition revealed by saccade trajectory modulation. J Neurophysiol 96: 1420 -1424. First published April 19, 2006 doi:10.1152/jn.00315.2006. 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. Suppressing saccades to distractors may additionally involve inhibiting corresponding 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 investigate 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 distractors. 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.
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.
In a series of experiments we examined the effects of the endogenous orienting of visual attention on human saccade latency. Three separate manipulations were performed: the orienting of visual attention, the prior offset of fixation (gap paradigm) and the bilateral presentation of saccade targets. Each of these manipulations was shown to make an independent contribution to saccade latency. In experiments 1 and 2 subjects were instructed to orient their attention covertly to a location by a verbal pre-cue; targets could appear in the attended hemifield (valid) or in the non-attended hemifield (invalid) together with a no-instruction (neutral) condition. Saccades were made under fixation gap and overlap conditions, to either single target or two bilaterally presented targets which appeared at equal and opposite eccentricities in both hemifields. The results showed a large increase (cost) of saccade latency to invalid targets and a small non-significant decrease (benefit) of saccade latency to valid targets. The cost associated with invalid targets replicates the "meridan crossing effect" shown in manual reaction time experiments and is consistent with the hemifield inhibition and premotor models of attentional orienting. The use of a "gap" procedure produced a generalised facilitation of saccade latency, which was not modified by the prior orienting of visual attention. The magnitude of the gap effect was similar for saccades made to attended and non-attended stimulis. This suggests that the gap effect may be due to ocular motor disengagement, or a warning signal effect, rather than to the prior disengagement of visual attention. When two targets were presented simultaneously, one in each hemifield, saccade latency was slowed compared with the single target condition. The magnitude of this slowing was unaffected by the prior orienting of visual attention or by the fixation condition. The slowing was examined in more detail in experiment 3, by presenting targets with brief offset delays. The latency increase was maximal if the two targets were presented simultaneously and decreased if the distractor appeared at short intervals (20-80 ms) before or after the saccade target onset. If the non-attended stimulus was presented at greater intervals (160, 240 ms) before the saccade target, then a facilitation effect was observed. This demonstrates that the onset of a distractor in the non-attended hemifield can have both an inhibitory and a facilitatory effect on a saccade production.
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