Spontaneous eye blinks are entrained by finger tapping.

We studied the mutual cross-talk between spontaneous eye blinks and continuous, self-paced unimanual and bimanual tapping. Both types of motor activities were analyzed with regard to their time-structure in synchronization-continuation tapping tasks which involved different task instructions, namely "standard" finger tapping (Experiment 1), "strong" tapping (Experiment 2) requiring more forceful finger movements, and "impulse-like" tapping (Experiment 3) where upward-downward finger movements had to be very fast. In a further control condition (Experiment 4), tapping was omitted altogether. The results revealed a prominent entrainment of spontaneous blink behavior by the manual tapping, with bimanual tapping being more effective than unimanual tapping, and with the "strong" and "impulse-like" tapping showing the largest effects on blink timing. Conversely, we found no significant effects of the tapping on the timing of the eye blinks across all experiments. The findings suggest a functional overlap of the motor control structures responsible for voluntary, rhythmic finger movements and eye blinking behavior.

Parietal damage dissociates saccade planning from presaccadic perceptual facilitation.

A well-known theory in the field of attention today is the premotor theory of attention which suggests that the mechanisms involved in eye movements are the same as those for spatial attention shifts. We tested a parietal damaged patient with unilateral optic ataxia and 4 controls on a dual saccade/attentional task and show a dissociation between saccadic eye movements and presaccadic perceptual enhancement at the saccade goal. Remarkably, though the patient was able to make the appropriate saccades to the left, impaired visual field (undistinguishable from saccades to his right, intact visual field), he was unable to discriminate the letter at the saccade goal (whereas his performance was like controls for letter discrimination in his right visual field). This suggests that saccade planning and presaccadic perceptual facilitation are separable--planning a saccade to a location does not necessitate that the processing of this location is enhanced. Based on these results, we suggest that the parietal cortex is necessary for the coupling between saccade planning and presaccadic perceptual facilitation.

Cerebellar lesions impair rapid saccade amplitude adaptation.

This study investigated whether the cerebellum is essential for rapid saccade adaptation. Saccade adaptation was elicited by 30% backward target steps during the primary saccade. Patients with cerebellar lesions adapted less than normal subjects, but saccade adaptation was most impaired in the group of patients with cerebellar degeneration. As the variability of the saccade gain in patients with cerebellar degeneration did not significantly differ from that in the other patients, the increased variability in motor performance due to a cerebellar lesion cannot alone explain this impaired adaptation.

Relative mislocalization of briefly presented stimuli in the retinal periphery.

We studied the ability to localize flashed stimuli, using a relative judgment task. When observers are asked to localize the peripheral position of a probe with respect to the midposition of a spatially extended comparison stimulus, they tend to judge the probe as being more toward the periphery than is the midposition of the comparison stimulus. We report seven experiments in which this novel phenomenon was explored. They reveal that the mislocalization occurs only when the probe and the comparison stimulus are presented in succession, independent of whether the probe or the comparison stimulus comes first (Experiment 1). The size of the mislocalization is dependent on the stimulus onset asynchrony (Experiment 2) and on the eccentricity of presentation (Experiment 3). In addition, the illusion also occurs in an absolute judgment task, which links mislocalization with the general tendency to judge peripherally presented stimuli as being more foveal than they actually are (Experiment 4). The last three experiments reveal that relative mislocalization is affected by the amount of spatial extension of the comparison stimulus (Experiment 5) and by its structure (Experiments 6 and 7). This pattern of results allows us to evaluate possible explanations of the illusion and to relate it to comparable tendencies observed in eye movement behavior. It is concluded that the system in charge of the guidance of saccadic eye movements is also the system that provides the metric in perceived visual space.

Visuomotor mental rotation of saccade direction.

This investigation studied the latencies of saccadic eye movements that were directed away from a target by a variable angular distance, which was given by instruction. Such a movement presumably requires an intentional, visuomotor mental rotation of the saccade vector, resulting in prolonged reaction times. From a study on the control of directed hand movements, it has been hypothesized that all visuomotor and visual mental rotation tasks share a common processing stage. We tested this hypothesis with a saccade task in which subjects shifted their gaze either towards (0 degrees, pro-saccade), or 30, 60, 90, 120, 150, or 180 degrees(anti-saccade) away from a randomly cued position on an imaginary clock face. With four different cueing conditions, latencies increased monotonically with required gaze shift from 0-150 degrees, thus exhibiting a mental rotation latency pattern. However, we also found anti-saccades faster than 150 degrees gaze shift and slower rotation speeds with peripheral cues than with central cues. Together with the overall shallower latency increase compared with previous findings with mental rotation tasks, these results cast doubt on the notion of a common, central processing mechanism for the different types of tasks.

Immediate post-saccadic information mediates space constancy.

We recently demonstrated that the perceived stability of a visual target that is displaced during a saccade critically depends on whether the target is present immediately when the saccade ends; blanking a target during and just after a saccade makes its intra-saccadic displacement more visible (Deubel et al. Vis Res 1996;36:985-996). Here, we investigate the interaction of visual context and blanking. Subjects saw a saccade target and an equal-sized distractor. During a saccade one or the other was displaced left or right. At the same time, one of the objects could be blanked briefly. Subjects reported whether the target or the distractor had jumped. The object that was blanked was more often seen as jumping (Experiment 1), regardless of which object really jumped, implying that continuously visible objects are preferentially perceived as stable. When both objects were blanked, longer blanking led to better accuracy at identifying which had jumped during a saccade. When one object was jumped and the other, stationary object was blanked (Experiment 2), the blanked object was mistakenly seen as jumping until the jump covered 50% or more of the saccade amplitude. In Experiment 3 a large continuously present texture underwent an undetected jump during a saccade, biasing judgments of simultaneous jumps of a blanked target. The results demonstrate that space constancy in normal situations is dominated by the assumption that a continuously present pattern is stable--this pattern becomes the spatial reference for the post-saccadic recalibration of perceptual space.

Effect of remote distractors on saccade programming: evidence for an extended fixation zone.

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.

P31 phosphor persistence at photopic mean luminance level.

P31 phosphor screens are frequently used for short-term presentation of dot and grating patterns, but experimental data obtained with this technique have been criticized because of possible parasitic effects of phosphor persistence on subjects visual performance. Recently, this issue provoked a controversial discussion in Vision Research (Groner et al., 1993; Westheimer, 1993, 1994; Irwin, 1994; Di Lollo et al., 1994) which was concerned with persistence effects of P31 screens for dot patterns. Supplementing this discussion, the present work deals with the effects of different types of patterns (dot pattern vs. gratings) and background mean-luminance levels (scotopic vs. phototopic) on phosphor persistence. Physical measurements of P31 persistence occurring with grating patterns of a mean luminance of 20 cd m-2 (i.e. photopic range) were obtained by using an extremely linear photometer with high temporal resolution. Under this photopic condition, the measurements demonstrate a fast decay of residual grating contrast to 1.4% of its original value within 50 ms after pattern offset. This phosphor behavior must be considered when designing an experiment with a P31 screen though it certainly embodies no problems in many applications.

Saccade target selection and object recognition: evidence for a common attentional mechanism.

The spatial interaction of visual attention and saccadic eye movements was investigated in a dual-task paradigm that required a target-directed saccade in combination with a letter discrimination task. Subjects had to saccade to locations within horizontal letter strings left and right of a central fixation cross. The performance in discriminating between the symbols "E" and "E", presented tachistoscopically before the saccade within the surrounding distractors was taken as a measure of visual attention. The data show that visual discrimination is best when discrimination stimulus and saccade target refer to the same object; discrimination at neighboring items is close to chance level. Also, it is not possible, in spite of prior knowledge of discrimination target position, to direct attention to the discrimination target while saccading to a spatially close saccade target. The data strongly argue for an obligatory and selective coupling of saccade programming and visual attention to one common target object. The results favor a model in which a single attentional mechanism selects objects for perceptual processing and recognition, and also provides the information necessary for motor action.

Postsaccadic target blanking prevents saccadic suppression of image displacement.

Displacement of a visual target during a saccadic eye movement is normally detected only at a high threshold, implying that high-quality information about target position is not stored in the nervous system across the saccade. We show that blanking the target for 50-300 msec after a saccade restores sensitivity to the displacement. With blanking, subjects reliably detect displacements as small as 0.33 deg across 6 deg eye movements, with correspondingly steep psychophysical functions. Performance with blanking in a fixation control is inferior, evidence for a saccadic enhancement of sensitivity to image displacement. If blanking is delayed so that the target is visible immediately after the saccade in its displaced position, performance declines to non-blanking levels. Blanking the target before the saccade, and restoring it during the saccade, yields a similar but weaker effect. We interpret these results with a model in which the visual system searches for the postsaccadic goal target within a restricted spatiotemporal window. If it is not found, the assumption of stationarity of the world is broken and the system makes use of other information such as extraretinal signals for calibrating location.

Rapid gain adaptation affects the dynamics of saccadic eye movements in humans.

The effect of rapid gain adaptation on the dynamics of visually guided saccades was investigated in six human subjects by using a search coil system. Saccadic adaptation was induced artificially by dislocating the target (by about 30% of the initial step) either forward (gain increase) or backward (gain decrease) during the primary saccade ("double-step paradigm"). Duration, peak velocity and peak acceleration and deceleration of a "standard 12 deg saccade" were computed from the data and were compared for the conditions of gain decrease, gain increase and the control without gain adaptation. The gain as well as the peak velocity and duration of the saccades showed an increased variability during the adaptation. In general, the abducting saccades had a higher peak acceleration than the adducting saccades, and all subjects showed an idiosyncratic pattern of the acceleration and deceleration. In the gain increase paradigm the subjects showed an increase in the duration and a decrease in the peak velocity. In the gain decrease paradigm there was a significant smaller ratio of peak acceleration/peak deceleration compared to the gain increase and the control condition. The findings demonstrate that rapid gain adaptation influences the dynamics of saccades in a specific way: peak saccadic velocity decreases and duration increases in the gain increase paradigm and peak acceleration/peak deceleration decreases in the gain decrease paradigm. Moreover, these results also suggest that the deceleration is neuronally controlled and not merely a result of mechanical constraints.

Separate adaptive mechanisms for the control of reactive and volitional saccadic eye movements.

Adaptive reduction of the gain of the saccadic system was induced by means of two basically different paradigms. In the first approach the subjects had to follow a step-wise moving target. During each follow-up saccade the target was systematically displaced by 25% of the initial step, into the opposite direction of the saccade. In the second approach the subjects scanned a display of six small items. During each scanning saccade the whole display was displaced by 25% into the opposite direction of the saccade. Both conditions lead to fast and consistent saccadic gain reductions. However, adaptation with the stepping target did not transfer to the saccades in the scanning situation, nor to delayed saccades in an overlap paradigm, nor to memory-guided saccades. Conversely, when saccades were adapted in the scanning situation, induced gain changes transferred to overlap and memory-guided saccades, but not to saccades following steps of a single target. The results suggest that two separate and largely independent mechanisms are involved in the generation of reactive, stimulus-triggered and volitional, internally generated saccades, respectively. Both types of responses can be selectively adapted.

Perceptual consequences of ocular lens overshoot during saccadic eye movements.

In a previous paper we compared eye globe records of saccadic eye movements (recorded with a scleral eye coil) with lens reflection records of the same eye movements (recorded with a dual-Purkinje-image eyetracker); we found evidence for considerable dynamic deviations between the two during and immediately after saccades. We ascribed these deviations to the movements of the eye's lens relative to the optical axis of the eye. This paper quantifies a predicted psychophysical effect of lens displacements during and after saccades. Two small targets, one above the other, were flashed for 2 msec in total darkness, the bottom one exactly at the end of the saccade, the top one 30 msec later. The first target appears deviated horizontally relative to the other, in a direction opposite to the saccade. Magnitude of the relative mislocalization can be up to 0.03 deg for each degree of saccadic eye movement. The result shows that the position of the visual image on the retina is affected both by position of the globe and by deviations of the lens from its normal location.

Fourth Purkinje image signals reveal eye-lens deviations and retinal image distortions during saccades.

Saccadic eye movements of various sizes and directions were registered simultaneously with the scleral search coil and a fifth-generation Dual Purkinje image eyetracker. Comparison of the search coil and the Purkinje image tracker records reveal considerable dynamic deviations during and immediately after the saccade, which we ascribe to the movements of the eye lens relative to the optical axis of the eye. Due to the increased stiffness of the tissues these deviations are smaller in older subjects. Also, they are larger at near accommodation. The size of the retinal image displacement which results from the lens movement proper can be as large as 0.5 deg, which may not be negligible in a number of visual tasks.

Reduced feedback frequency enhances generalized motor program learning but not parameterization learning.

The purpose of the study was to examine the effects of a reduced feedback frequency on the learning of generalized motor programs and movement parameterization. Subjects practiced three movement patterns with the same relative timing and the same relative amplitude, but with varied movement time (Experiment 1) or varied movement amplitude (Experiment 2). KR was given either on 100% or 63% of the trials, with learning being assessed by retention and transfer tests. In both experiments, reduced KR frequency enhanced GMP learning but generally degraded parameter learning. These data provide converging evidence for the dissociation of the program and parameterization processes postulated in GMP theory.

Sensory and motor aspects of saccade control.

The oculomotor reactions have recently attracted increasing attention for diagnostic purposes. This is in line with the view that the oculomotor system is one of the simpler, machine-like sensorimotor systems. This paper presents two examples to demonstrate that the complexity of sensorimotor processing may be higher than expected from the outcome of experiments under rather restricted stimulus conditions. In the first part it is shown that complex preprocessing including the evaluation of spatial gradients of visual structure is an integral part of the programming of reflex-like saccades. The second part concerns adaptivity of saccadic eye movements. The data demonstrate that saccadic gain control is highly specific to the direction of the saccade. It is suggested that many central deficits may be hidden as a consequence of the effect of specific adaptive mechanisms.

Saccadic eye movements and the detection of fast-moving gratings.

Experiments are presented in which the effect of saccadic eye movements on the visibility of sinusoidal gratings drifting with velocities between 2 deg/s and 400 deg/s is investigated. The results demonstrate that saccades are highly useful for detecting this class of stimuli. Due to a saccade, otherwise subthreshold stimuli become visible as short, distinct flashes of the seemingly stationary pattern. The paper analyzes in detail the dependence of the amount of facilitation on saccade size and relative direction and isolates the additional effect of saccadic suppression. A simple model is proposed which predicts the experimental findings.

Adaptive gain control of saccadic eye movements.

Properties of gain adaptivity in the saccadic system were studied. Subjects had to track a target which moved in single or double steps. The first target step which elicited the primary saccade had an amplitude in the range of 8-16 deg. The primary saccade triggered a further target displacement of 4 deg either in the same or--in different experimental sessions--in the opposite direction of the first target step. These consistent intrasaccadic target displacements lead to adaptive changes of saccadic amplitudes. The experimental data show that the saccadic system adapts to the stimulus sequence in a simple, parametric manner, namely by changing its gain. Consequently, it is assumed that a single gain element determines saccade sizes for all target eccentricities. Further, it is shown that adaptation has different time courses for gain increase and decrease, and its performance is close to completeness. The results are discussed with respect to the undershooting behaviour of goal-directed saccades and the functional demands to the saccadic system.

Threshold perception and saccadic eye movements.

Involuntary eye movements were recorded during threshold detection tasks under various experimental conditions. The data were analyzed for interdependencies between stimulus parameters, detection performance, and oculomotor behaviour. The data demonstrate that under certain conditions, saccadic parameters are adaptive to specific stimulus properties. Further, the data suggest that for stationary patterns with low spatial frequencies and for gratings flickering with high temporal frequencies, detection is facilitated considerably by the occurrence of a saccadic eye movement. These facilitation effects are consistent with the predictions of a theoretical model presented in a previous paper.

The effect of saccades on threshold perception--a model study.

The effect of saccadic eye movements on threshold perception is investigated theoretically. The proposed model considers eye movements by taking into account the shifting of the stimulus pattern on the retina during the occurrence of an eye movement. Saccades are characterized by high velocity and short duration. These motions cause overshoots in the response of linear filters to certain stimulus patterns. Therefore, the model predicts facilitation effects of saccades in the perception of low spatial frequency patterns and patterns flickering with high temporal frequencies. These results agree with experimentally obtained data presented in a subsequent paper. A simple approach is formulated which approximates the complex shifting function of a saccade by a switching of the pattern.