Localisation of auditory targets during optokinetic nystagmus

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.     

Auditory localization: The importance of eye movements and a textured visual environment

Two experiments were designed to investigate the factors involved in the visual facilitation of auditory localization. In both experiments, adult human Ss pointed to targets in a variety of visual conditions. The results of the first experiment showed that target-directed eye movements were important. In the second experiment, eye localization was assessed, along with pointing localization. Both eye and hand localization of the hidden auditory targets were better when target-directed eye movements were made in a lighted environment than when made in the dark. Data also suggested that Ss have better knowledge of their eye position in the light. Possible mechanisms for the involvement of eye movements were suggested, and the theoretical importance of the results was discussed.     

Programming saccadic eye movements

This article addresses questions about the preparatory processes that immediately precede saccadic eye movements. Saccade latencies were measured in a task in which subjects were provided partial advance information about the spatial location of a target fixation. In one experiment, subjects were faster in initiating saccades when they knew either the direction or amplitude of the required movement in advance compared to a condition with equal uncertainty about the number of potential saccade targets but without knowledge of the parameters required to execute the movement. These results suggest that the direction and amplitude for an upcoming saccade were calculated separately, and not in a fixed serial order. In another experiment, subjects appear to have programmed the saccades more holistically--with computations of direction and amplitude parameters occurring simultaneously. The implications of these results for models of eye movement preparation are discussed.     

Functional between-hand differences and outflow eye position information

Pointing accuracy with an unseen hand to a just-extinguished visual target was examined in various eye movement conditions. When subjects caught the target by a saccade, they showed about the same degree of accuracy as that shown in pointing to a visible target. On the other hand, when subjects tracked a moving target by a pursuit eye movement, they systematically undershot when subsequently pointing to the target. The differential effect of the two types of eye movements on pointing tasks was examined on both the preferred and non-preferred hands, and it was found that the effect of eye movements was more prominent on the preferred hand than on the non-preferred hand. The results are discussed in relation to outflow eye position information.     

Memory representations guide targeting eye movements in a natural task

The change blindness phenomenon suggests that visual representations retained across saccades are very limited. In this paper we sought to specify the kind of information that is in fact retained. We investigated targeting performance for saccadic eye movements, since one need for visual representations across eye and body positions may be to guide coordinated movements. We examined saccades in the context of an ongoing sensory motor task in order to make stronger generalizations about natural visual functioning and deployment of attention. Human subjects copied random patterns of coloured blocks on a computer display. Their eye movement pattern was consistent from block to block, including a precise saccade to a previously-placed, neighbouring block during each additional block placement. This natural, consistent eye movement allowed the previously-placed, neighbouring block to serve as an implicit target without instructions to the subject. On random trials, we removed the target object from the display during a preceding saccade, so that observers were required to make the targeting saccade without a currently visible target. Targeting performance was excellent, and appeared to be influenced by spatial information that was not visible during the preceding fixation. Subjects were generally unaware of the disappearance and reappearance of the target. We conclude that spatial information about visual targets is retained across eye movements and used to guide subsequent movements.     

Retinal and extraretinal information in movement perception: how to invert the Filehne illusion

During a pursuit eye movement made in darkness across a small stationary stimulus, the stimulus is perceived as moving in the opposite direction to the eyes. This so-called Filehne illusion is usually explained by assuming that during pursuit eye movements the extraretinal signal (which informs the visual system about eye velocity so that retinal image motion can be interpreted) falls short. A study is reported in which the concept of an extraretinal signal is replaced by the concept of a reference signal, which serves to inform the visual system about the velocity of the retinae in space. Reference signals are evoked in response to eye movements, but also in response to any stimulation that may yield a sensation of self-motion, because during self-motion the retinae also move in space. Optokinetic stimulation should therefore affect reference signal size. To test this prediction the Filehne illusion was investigated with stimuli of different optokinetic potentials. As predicted, with briefly presented stimuli (no optokinetic potential) the usual illusion always occurred. With longer stimulus presentation times the magnitude of the illusion was reduced when the spatial frequency of the stimulus was reduced (increased optokinetic potential). At very low spatial frequencies (strongest optokinetic potential) the illusion was inverted. The significance of the conclusion, that reference signal size increases with increasing optokinetic stimulus potential, is discussed. It appears to explain many visual illusions, such as the movement aftereffect and center-surround induced motion, and it may bridge the gap between direct Gibsonian and indirect inferential theories of motion perception.     

Effects of eye movements on the recognition and localization of dichotic stimuli

Three experiments examined the effect of gaze shifts on overall performance and ear differences in dichotic listening. In the first two experiments, lights were switched on and off so as to induce rightward, leftward, or upward gaze during dichotic stimulation. The dichotic material consisted of musical passages in Experiment 1 and two kinds of verbal material in Experiment 2. Vertical eye movements enhanced the accuracy of identification of music but not verbal material. The lateral direction of eye movements affected subjects' ability to localize targets in Experiment 1: localization was more accurate in the direction toward which subjects were looking. In the third experiment it was found that optokinetic nystagmus (OKN) influenced the asymmetry of performance on a dichotic consonant-vowel (CV) test. The right-ear advantage was greatest when the OKN drum rotated from left to right and least when it rotated from right to left. The effect was due to corresponding variation in left-ear scores. Possible mechanisms through which shifts of gaze affect auditory identification and localization are proposed.     

Two kinds of adaptation in the constancy of visual direction and their different effects on the perception of shape and visual direction

Adaptation in the constancy of visual direction can be obtained under two radically different conditions, called eye-movement adaptation and field adaptation. Adaptation resulting from these conditions and from a “normal” condition was measured with a newly developed estimation test. Eye-movement adaptation was found to cause an alteration of compensatory eye movements. It apparently consists of a changed evaluation of eye movements, as demonstrated by two different pointing tests. A form test where the shape of a large oblong is set to look square also confirmed this interpretation. After field adaptation, a pointing test did not register a change, but an adaptation effect could be measured with a forward direction test. This test and a square test where no eye movements were permitted proved to be specific to field adaptation; they measured no effect after eye-movementadaptation. The normal adaptation condition Was apparently equivalent to the eye-movement adaptation condition. Its effect could be measured only with a pointing test. When we changed the normal adaptation condition so that frequent saccades were made during head turning, strong effects were measured with the two tests that were specific to field adaptation.     

Adaptation to visual and proprioceptive rearrangement: Origin of the differential effectiveness of active and passive movements

In Experiment 1, subjects exposed to a discordance between the visual and ”proprioceptive” locations of external targets were found to exhibit aftereffects when later pointing without sight of their hands at visual targets. Aftereffects occur both when the discordance is introduced in the traditional fashion by displacing the visual locations of targets and when the proprioceptive locations of targets are displaced. These observations indicate that there is nothing unique about the visual rearrangement paradigm—the crucial factor determining whether adaptation will be elicited is the presence of a discordance in the positional information being conveyed over two different sensory modalities. In a second experiment, the effectiveness of active and passive movements in eliciting adaptation was studied using an experimental paradigm in which subjects were exposed to a systematic discordance between the visual and proprioceptive locations of external targets without ever being permitted sight of their hands; a superiority of active movements was observed, just as is usually found in visual rearrangement experiments in which sight of the hand is permitted. Evidence is presented that the failure of passive movements to elicit adaptation is related to a deterioration in accuracy of position sense information during passive limb movement.     

Induced motion of a fixated target: influence of voluntary eye deviation.

Induced motion (IM) was observed in a fixated target in the direction opposite to the real motion of a moving background. Relative to a fixation target located straight ahead, IM decreased when fixation was deviated 10 degrees in the same direction as background motion and increased when fixation was deviated 10 degrees opposite background motion. These results are consistent with a "nystagmus-suppression" hypothesis for subjective motion of fixated targets: the magnitude of illusory motion is correlated with the amount of voluntary efference required to oppose involuntary eye movements that would occur in the absence of fixation. In addition to the form of IM studied, this explanation applies to autokinesis, apparent concomitant motion, and the oculogyral illusion. Accounts of IM that stress visual capture of vection, afferent mechanisms, egocenter deviations, or phenomenological principles, although they may explain some forms of IM, do not account for the present results.     

An asymmetry in the control of eye movements and shifts of attention

Under conditions of monocular viewing, eye movements tend to be directed toward stimuli in the temporal visual field. The present experiments indicate that this bias does not extend to shifts of attention under conditions in which the eyes remain fixed. Experiments 1 and 2 support this conclusion for voluntary shifts of attention, experiment 3 for involuntary shifts of attention. These results support Posner and Cohen's (1980) hypothesis that the temporal bias reflects the properties of an isolable component of the eye movement control system and extend previous dissociations of the mechanisms underlying the control of shifts of attention and eye movements.     

Masking of foveal and parafoveal vision during eye fixations in reading

A window or visual mask as moved across text in synchrony with the reader's eye movements. The size of the window or mask was varied so that either information in foveal or parafoveal vision was masked on each fixation. In another experiment, the onset of the mask was delayed for a certain amount of time following the end of the saccade. The results of the experiments point out the relative importance of foveal and parafoveal vision for reading and further indicate that most of the visual information necessary for reading can be acquired during the first 50 msec that information is available during an eye fixation.     

Smooth pursuit eye movements in normal and dyslexic children

This paper describes a detailed study of horizontal eye movements associated with visual tracking of a smoothly moving target. Essentially all children, even at target velocities as low as 5 degrees/sec., show some saccadic eye movements superimposed on smooth tracking movements. Detailed analysis of pursuit eye-movements from a group of 26 poor readers and 34 normal controls (8 to 13 yr.) showed that about 25% of poor readers have an abnormally raised saccadic component in smooth pursuit. This suggests that studies of eye movements during tracking of smoothly moving targets at low velocity, combined with a quantitative approach to data analysis, may be useful for early detection of a significant proportion of poor-reading children.     

Distractor interference during smooth pursuit eye movements

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.     

Induced motion of a fixated target: Influence of voluntary eye deviation

Induced motion (IM) was observed in a fixated target in the direction opposite to the real motion of a moving background. Relative to a fixation target located straight ahead, IM decreased when fixation was deviated 10° in the same direction as background motion and increased when fixation was deviated 10° opposite background motion. These results are consistent with a “nystagmus-suppression” hypothesis for subjective motion of fixated targets: the magnitude of illusory motion is correlated with the amount of voluntary efference required to oppose involuntary eye movements that would occur in the absence of fixation. In addition to the form of IM studied, this explanation applies to autokinesis, apparent concomitant motion, and the oculogyral illusion. Accounts of IM that stress visual capture of vection, afferent mechanisms, egocenter deviations, or phenomenological principles, although they may explain some forms of IM, do not account for the present results.     

Structured perceptual arrays and the modulation of Fitts's law: examining saccadic eye movements

On the basis of recent observations of a modulation of Fitts's law for manual pointing movements in structured visual arrays (J. J. Adam, R. Mol, J. Pratt, & M. H. Fischer, 2006; J. Pratt, J. J. Adam, & M. H. Fischer, 2007), the authors examined whether a similar modulation occurs for saccadic eye movements. Healthy participants (N = 19) made horizontal saccades to targets that appeared randomly in 1 of 4 positions, either on an empty background or within 1 of 4 placeholder boxes. Whereas in previous studies, placeholders caused a decrease in movement time (MT) without the normal decrease in movement accuracy predicted by Fitts's law, placeholders in the present experiment increased saccadic accuracy (decreased endpoint variability) without an increase in MT. The present results extend the findings of J. J. Adam et al. of a modulation of Fitts's law from the temporal domain to the spatial domain and from manual movements to eye movements.     

Visual attention amplifies response priming of pointing movements to color targets

We studied the influence of spatial visual attention on the time course of primed pointing movements. We measured pointing responses to color targets preceded by color stimuli priming either the same response as the target or the opposite response. The effects of visual attention at the prime and target locations were studied by varying both the cue-prime and prime-target intervals when presenting either exogenous attentional cues or, in a separate experiment, endogenous cues whose processing was a precondition for performing the task. Pointing trajectories revealed large priming effects in which pointing responses were first controlled by prime signals and then captured in midflight by target signals. Priming effects were strongly amplified when the relevant prime locations were visually attended at optimal cue-prime SOAs, with attention modulating the entire time course of the primed pointing movements. We propose that visual attention amplifies the earliest waves of visuomotor feedforward information, elicited in turn by primes and by targets.     

Covert shifts of attention precede involuntary eye movements

There is considerable evidence that covert visual attention precedes voluntary eye movements to an intended location. What happens to covert attention when an involuntary saccadic eye movement is made? In agreement with other researchers, we found that attention and voluntary eye movements are tightly coupled in such a way that attention always shifts to the intended location before the eyes begin to move. However, we found that when an involuntary eye movement is made, attention first precedes the eyes to the unintended location and then switches to the intended location, with the eyes following this pattern a short time later. These results support the notion that attention and saccade programming are tightly coupled.     

Eye movement indices of mental workload

Four investigations were carried out to assess the feasibility of using eye movement measures as indices of mental workload. In the first experiment, saccadic extent was measured during free viewing while subjects performed low, moderate and high complexity, auditory tone counting as the workload tasks. The range of saccadic extent decreased significantly as tone counting complexity (workload) was increased. In the second experiment the range of spontaneous saccades was measured under three levels of counting complexity with a visual task that did not require fixation or tracking. The results indicated that the extent of saccadic eye movements was significantly restricted as counting complexity increased. In the third experiment, the effects of practice were examined and decreased saccadic range under high tone counting complexity was observed even when significant increases in performance occurred with practice. Finally, in experiment 4, the first experiment was repeated with additional optokinetic stimulation and the saccadic range was again observed to decrease with tone counting complexity. The results indicated that the extent of spontaneous and elicited eye movements was significantly restricted as counting complexity increased. We conclude that this measure may provide a valuable index of mental workload.     

An investigation of the relationship between eye and retinal image movement in the perception of movement

The question investigated was whether or not eye movements accompanied by abnormal retinal image movements, movements that are either or both at a different rate or in a different direction than the eye movement, predictably lead to perceived movement. Os reported whether or not they saw a visual target move when the movement of the target was either dependent on and simultaneous with their eye movements or when the target movement was independent of their eye movements. In the main experiment, observations were made when the ratio between eye and target movement fem/tm) was 2/5, 1/5, 1/10, 1/20, and 0. All these ratios were tested when the direction of the target movement was in the same (H+), opposite (H?), and at right angles to (V+, V?) the movement of the eyes. Eye movements, target movements, and reports of target movement were recorded. Results indicate that a discrepancy between eye and target movement greater than 20% predictably leads to perceived target movement, whereas a discrepancy of 5% or less rarely leads to perceived movement. The results are interpreted as support for the operation of a compensatory mechanism during eye movements.