Oculomotor Readiness Research Articles

Is covert spatial orienting embodied or disembodied cognition? A historical review.

The possible relations between eye movements and shifts of attention are considered in the context of the contemporary proposal of embodied cognition. The focus of this historical review is Klein's oculomotor readiness hypothesis for how visuospatial attention might be allocated when under endogenous control. When eye movements are actually executed, attention shifts in advance of these movements. But when eye movements are prepared but not executed, shifts of attention are not observed. Conversely, when attention is allocated endogenously and covertly to a location in space, eye movements to that location are not prepared. These findings suggest that covert spatial orienting when under endogenous control is more dis-embodied than embodied cognition.

Associations and Dissociations between Oculomotor Readiness and Covert Attention.

The idea that covert mental processes such as spatial attention are fundamentally dependent on systems that control overt movements of the eyes has had a profound influence on theoretical models of spatial attention. However, theories such as Klein’s Oculomotor Readiness Hypothesis (OMRH) and Rizzolatti’s Premotor Theory have not gone unchallenged. We previously argued that although OMRH/Premotor theory is inadequate to explain pre-saccadic attention and endogenous covert orienting, it may still be tenable as a theory of exogenous covert orienting. In this article we briefly reiterate the key lines of argument for and against OMRH/Premotor theory, then evaluate the Oculomotor Readiness account of Exogenous Orienting (OREO) with respect to more recent empirical data. These studies broadly confirm the importance of oculomotor preparation for covert, exogenous attention. We explain this relationship in terms of reciprocal links between parietal ‘priority maps’ and the midbrain oculomotor centres that translate priority-related activation into potential saccade endpoints. We conclude that the OMRH/Premotor theory hypothesis is false for covert, endogenous orienting but remains tenable as an explanation for covert, exogenous orienting.

Does oculomotor readiness mediate exogenous capture of visual attention?

The oculomotor readiness hypothesis makes 2 predictions: Shifts in covert attention are accompanied by preparedness to move one's eyes to the attended region, and preparedness to move one's eyes to a region in space is accompanied by a shift in covert attention to the prepared location. Both predictions have been disconfirmed using an endogenous attention task. In the 2 experiments presented here, the same 2 predictions were tested using an exogenous attention task. It was found that participants experienced covert capture without accompanying oculomotor activation and experienced oculomotor activation without accompanying covert capture. While under everyday conditions the overt and covert orienting systems may be strongly linked, apparently they can nonetheless operate with a high degree of independence from one another. (PsycINFO Database Record

Motor output evoked by subsaccadic stimulation of primate frontal eye fields

In addition to its role in shifting the line of sight, the oculomotor system is also involved in the covert orienting of visuospatial attention. Causal evidence supporting this premotor theory of attention, or oculomotor readiness hypothesis, comes from the effect of subsaccadic threshold stimulation of the oculomotor system on behavior and neural activity in the absence of evoked saccades, which parallels the effects of covert attention. Here, by recording neck-muscle activity from monkeys and systematically titrating the level of stimulation current delivered to the frontal eye fields (FEF), we show that such subsaccadic stimulation is not divorced from immediate motor output but instead evokes neck-muscle responses at latencies that approach the minimal conduction time to the motor periphery. On average, neck-muscle thresholds were approximately 25% lower than saccade thresholds, and this difference is larger for FEF sites associated with progressively larger saccades. Importantly, we commonly observed lower neck-muscle thresholds even at sites evoking saccades <or=5 degrees in magnitude, although such small saccades are not associated with head motion. Neck-muscle thresholds compare well with the current levels used in previous studies to influence behavior or neural activity through activation of FEF neurons feeding back to extrastriate cortex. Our results complement this previous work by suggesting that the neurobiologic substrate that covertly orients visuospatial attention shares this command with head premotor circuits in the brainstem, culminating with recruitment in the motor periphery.

Attentional release in the saccadic gap effect

Can a release of attention from fixation help explain the saccadic ‘gap effect’, the shortening of saccadic latency (SL) when the fixation spot is extinguished just before saccade target onset? Practiced observers generated SLs and button-presses to one of four 10° eccentric targets in overlap (fixation spot stays on), gap0 (fixation offsets at target onset), and gap200 conditions; in gap200, the fixation spot was removed, dimmed, expanded, or brightened 200 ms before target onset. Our data excluded speed-accuracy trade-offs, express saccades, stimulus salience, and oculomotor readiness, while fixation offset and general warning had minor effects, leaving attention release as the default explanation. Supporting this notion, finger-press reactions to foveal probe dots presented after the fixation spot was brightened (to hold attention) were faster than those made after the spot was removed (to release attention). Varying the time from gap onset to the probe dot mapped out the time-course of the putative attentional release, which takes ∼140 ms.

Control of oculomotor reflexes: independent effects of strategic and automatic preparation

The reduction in saccade latency when the fixation point is removed (fixation offset effect-FOE) reflects the degree to which fixation neurons are under influence by a stimulus at fixation. Strategic manipulations of oculomotor readiness that bring these neurons under endogenous control reduce the magnitude of the FOE. Using an aging foreperiod paradigm, and the FOE as a marker for cortical control of reflexive fixation, we showed that, for both prosaccades and antisaccades, increasing preparation across the foreperiod reduced both saccade latency and the FOE. Consistent with Los's trace conditioning account, these effects reflected greater preparation for trials when the current short foreperiod was preceded by a trial with a short foreperiod. The FOE was also smaller for antisaccades than for prosaccades demonstrating strategic modulation. However, the effects of trace conditioning were comparable in the two tasks, demonstrating that strategic and unconscious priming effects both independently modulate the control of ocular fixation.

Role of Visual Attention in Cognitive Control of Oculomotor Readiness in Students with Reading Disabilities

This study investigated eye movement and comprehension therapy in Grade 6 children with reading disabilities (RD). Both order of therapy and type of therapy were examined. Furthermore, the implications of visual attention in ameliorating reading disability are discussed. Thirty-one students with RD were identified using standardized reading comprehension tests. Eye movements were analyzed objectively using an infra-red recording device. Reading scores of participating children were 0.5 to 1 SD below the national mean. Testing took place before the start of therapy (T1) and was repeated after 12 weeks (T2) and 24 weeks (T3) of therapy. One group of students had eye movement therapy first, followed by comprehension therapy; in the other group, the order was reversed. Data were evaluated using a repeated measures MANOVA and post hoc tests. At T1, mean reading grade was 2 years below grade level, and eye movement scores were at about Grade 2 level. Mean growth in reading comprehension for the total sample was 2.6 years (p < .01) at T3; equally significant improvement was measured in eye movements (p < .01). Learning rate in reading comprehension improved from 60% (T1) to 400% (T3). Although within-group differences were statistically significant, between-group differences were not significant for comprehension or eye movements. Order of therapy (comprehension first or eye movements first) was not significant. Improvements in within-group scores for comprehension and eye movements were consistently significant at T2 and T3. Eye movement therapy improved eye movements and also resulted in significant gains in reading comprehension. Comprehension therapy likewise produced improvement both in eye movement efficiency and in reading comprehension. The results support the notion of a cognitive link among visual attention, oculomotor readiness, and reading comprehension.

Spatial attention and latencies of saccadic eye movements

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

Oculomotor readiness and covert orienting: differences between central and peripheral precues.

The costs produced by invalid precues can depend on the spatial relationship between the cued location and the target location. If oculomotor programs mediate attention shifts, then the effect of varying the spatial relation between the cue and target should be the same for covert orienting (indexed by manual responses) and saccadic responses. We found this to be true only for central precues. With central precues, both manual and saccadic costs were greater when cue and target occurred on opposite sides of the vertical meridian than when they occurred on the same side. With peripheral precues, there were no meridian effects in either response condition, but there was a significant dissociation in the pattern of saccadic and manual costs. For manual responses costs were greater when the target was eccentric relative to the cue, whereas for saccades costs were greater when the cue was eccentric to the target. These results provide additional support for the idea that different orienting mechanisms are engaged by central and peripheral precues. They further suggest that the relationship between oculomotor and attentional orienting may depend on the nature of the precue, with the potential for interdependence being greater with central precues.

The reduction of saccadic latency by prior offset of the fixation point: An analysis of the gap effect

The latency to initiate a saccade (saccadic reaction time) to an eccentric target is reduced by extinguishing the fixation stimulus prior to the target onset. Various accounts have attributed this latency reduction (referred to as the gap effect) to facilitated sensory processing, oculomotor readiness, or attentional processes. Two experiments were performed to explore the relative contributions of these factors to the gap effect. Experiment 1 demonstrates that the reduction in saccadic reaction time (RT) produced by fixation point offset is additive with the effect of target luminance. Experiment 2 indicates that the gap effect is specific for saccades directed toward a peripheral target and does not influence saccades directed away from the target (i.e., antisaccades) or choice-manual RT. The results are consistent with an interpretation of the gap effect in terms of facilitated premotor processing in the superior colliculus.