Control' of reflexive and voluntary saccades in the gap effect.

In two experiments, we examined whether voluntary and reflexive saccades shared a common fixation disengagement mechanism. Participants were required to perform a variety of tasks, each requiring a different level of information processing of the display prior to execution of the saccade. In Experiment 1, participants executed either a prosaccade or an antisaccade upon detecting a stimulus array. In Experiment 2, participants executed a prosaccade to a stimulus array only if the array contained a target item. The target could be a line (easy search) or a digit (difficult search). The critical manipulation in both experiments was the relative timing between the removal of the fixation stimulus and the onset of the stimulus array. In both experiments, it was found that saccadic latencies were shortest when the fixation stimulus was removed before the onset of the stimulus array--a gap effect. It was concluded that reflexive and voluntary saccades share a common fixation disengagement mechanism that is largely independent of higher level cognitive processes.     

Strategic control over saccadic eye movements: studies of the fixation offset effect

We studied the strategic (presumably cortical) control of ocular fixation in experiments that measured the fixation offset effect (FOE) while manipulating readiness to make reflexive or voluntary eye movements. The visual grasp reflex, which generates reflexive saccades to peripheral visual signals, reflects an opponent process in the superior colliculus (SC) between fixation cells at the rostral pole, whose activity helps maintain ocular position and increases when a stimulus is present at fixation, and movement cells, which generate saccades and are inhibited by rostral fixation neurons. Voluntary eye movements are controlled by movement and fixation cells in the frontal eye field (FEF). The FOE--a decrease in saccade latency when the fixation stimulus is extinguished--has been shown to reflect activity in the collicular eye movement circuitry and also to have an activity correlate in the FEF. Our manipulation of preparatory set to make reflexive or voluntary eye movements showed that when reflexive saccades were frequent and voluntary saccades were infrequent, the FOE was attenuated only for reflexive saccades. When voluntary saccades were frequent and reflexive saccades were infrequent, the FOE was attenuated only for voluntary saccades. We conclude that cortical processes related to task strategy are able to decrease fixation neuron activity even in the presence of a fixation stimulus, resulting in a smaller FOE. The dissociation in the effects of a fixation stimulus on reflexive and voluntary saccade latencies under the same strategic set suggests that the FOEs for these two types of eye movements may reflect a change in cellular activity in different neural structures, perhaps in the SC for reflexive saccades and in the FEF for voluntary saccades.     

反向眼跳的实验范式、机制及影响因素

反向眼跳任务是研究内源性眼跳的主要方法。1978年,Hallett在研究中首次使用了反向眼跳的实验任务。实验中要求被试抑制对外围目标的注视,并注视它的相反位置。反向眼跳任务是研究行为控制以及注意功能的有效范式。目前认为反向眼跳任务中的朝向眼跳和反向眼跳计划是同时加工并相互竞争的,并且反向眼跳的产生与额顶叶皮层下组织有关。反向眼跳会受到多种因素的影响,例如,空白效应、工作记忆、认知老化、目标离心率等。     

Presaccadic attention allocation and express saccades

Express saccades are visually-guided saccades that are characterized by an extremely short latency of about 100 ms. The present experiments tested the hypothesis that a disengagement of visual attention is necessary for the generation of express saccades. All subjects produced large numbers of express saccades in the gap paradigm, in which the fixation stimulus is removed 200 ms before target onset (Exp. 1), but not in the overlap paradigm, in which the fixation stimulus remains on during the entire trial (Exp. 2). By means of peripheral cues (Exps. 3–5) and central cues (Exps. 6–7), visual attention was directed at the target location for the saccade before the actual appearance of the saccade target. In all experiments, the location cues facilitated rather than abolished express saccades. The generation of express saccades was facilitated even when the currently fixated visual stimulus was not removed before target onset (fixation-overlap; Exps. 5–7). The results are explained by the hypothesis that a disengagement of a separate fixation system is necessary for the generation of express saccades, a hypothesis that is in line with current neurobiological findings.     

A controlled-attention view of working-memory capacity.

In 2 experiments the authors examined whether individual differences in working-memory (WM) capacity are related to attentional control. Experiment 1 tested high- and low-WM-span (high-span and low-span) participants in a prosaccade task, in which a visual cue appeared in the same location as a subsequent to-be-identified target letter, and in an antisaccade task, in which a target appeared opposite the cued location. Span groups identified targets equally well in the prosaccade task, reflecting equivalence in automatic orienting. However, low-span participants were slower and less accurate than high-span participants in the antisaccade task, reflecting differences in attentional control. Experiment 2 measured eye movements across a long antisaccade session. Low-span participants made slower and more erroneous saccades than did high-span participants. In both experiments, low-span participants performed poorly when task switching from antisaccade to prosaccade blocks. The findings support a controlled-attention view of WM capacity.     

Fixation disengagement and eye-movement latency

We examined eye-movement latencies to a target that appeared during visual fixation of a stationary stimulus, a moving stimulus, or an extrafoveal stimulus. The stimulus at fixation was turned off either before target onset (gap condition) or after target onset (overlap condition). Consistent with previous research, saccadic latencies were shorter in gap conditions than they were in overlap conditions (the gap effect). In Experiment 1, a gap effect was observed for vergence eye movements. In Experiment 2, a gap effect was observed for saccades directed at a target that appeared during visual pursuit of a moving stimulus. In Experiment 3, a gap effect was observed for saccades directed at a target that appeared during extrafoveal fixation. The present results extend reports of the gap effect for saccadic shifts during visual fixation to (a) vergence shifts during visual fixation, (b) saccadic shifts during smooth visual pursuit, and (c) saccadic shifts during extrafoveal fixation. The present findings are discussed with respect to the incompatible goals of fixation-locking and fixation-shifting oculomotor responses.     

Saccade landing point selection and the competition account of pro- and antisaccade generation: the involvement of visual attention--a review

This paper presents a review and summary of experimental findings on the role of attention in the preparation of saccadic eye movements. The focus is on experiments where performance of prosaccades (saccades towards a suddenly appearing item) and antisaccades (saccades of equal amplitude in the direction opposite to where the target moved) is compared. Evidence suggests that these two opposite responses to the same stimulus event entail competition between neural pathways that generate reflexive movements to the target and neural mechanisms involved in inhibiting the reflex and generating a voluntary gaze shift in the opposite direction to the target appearance. Evidence for such a competition account is discussed in light of a large amount of experimental findings and the overall picture clearly indicates that this competition account has great explanatory power when data on saccadic reaction times and error rates are compared for the two types of saccade. The role of attention is also discussed in particular in light of the finding that the withdrawal of attention by a secondary task 200 to 500 ms before the saccade target appears, leads to speeded antisaccades (without a similar increase in error rates), showing that the results do not simply reflect a speed-accuracy trade-off. This result indicates that the tendency for "reflexive" prosaccades is diminished when attention is engaged in a different task. Furthermore, experiments are discussed that show that as the tendency for a reflexive prosaccade is weakened, antisaccades are speeded up, further supporting the competition account of pro- and antisaccade generation. In the light of evidence from neurophysiology of monkeys and humans, a tentative model of pro- and antisaccade generation is proposed.     

注意控制定势和线索类型在注意捕获中的作用

采用前线索范式, 通过3个实验考察了注意控制定势和线索类型在注意捕获中的作用。实验1要求被试搜索颜色奇异项目标或突现目标, 涉及较弱的注意控制定势, 结果表明, 突现线索在作为匹配线索和不匹配线索时都能捕获注意, 而颜色奇异项线索只有在作为匹配线索时才能捕获注意。实验2要求被试搜索具体特征的红色或白色目标, 使被试建立较强的特征搜索定势, 结果表明, 颜色奇异项线索和突现线索在作为匹配线索时都能捕获注意, 但是在不匹配条件下, 颜色奇异项线索被抑制了, 而突现线索不能捕获注意; 实验3采取go/no-go范式, 要求被试对红色目标进行反应, 对白色目标进行抑制, 并通过设置不同的线索-目标时间间隔来进一步考察注意捕获的机制, 结果表明, 在600 ms间隔下, no-go突现线索被抑制, 而对于go颜色奇异项线索来说, 随着时间间隔的增加, 注意捕获效应减少。结果与相倚捕获假说和抑制独立捕获假说不符, 支持注意转移假说：自上而下的注意控制定势通过把注意从和目标不匹配的特征位置转移发挥作用; 颜色奇异项线索和突现线索的注意捕获机制相同, 但是突现线索相对于颜色奇异项线索来说, 在与目标不匹配的条件下, 更容易捕获注意, 更难被抑制。     

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.     

The effects of eye and limb movements on working memory

Three experiments examined the role of eye and limb movements in the maintenance of information in spatial working memory. In Experiment 1, reflexive saccades interfered with memory span for spatial locations but did not interfere with memory span for letters. In Experiment 2, three different types of eye movements (reflexive saccades, pro-saccades, and anti-saccades) interfered with working memory to the same extent. In all three cases, spatial working memory was much more affected than verbal working memory. The results of these two experiments suggest that eye movements interfere with spatial working memory primarily by disrupting processes localised in the visuospatial sketchpad. In Experiment 3, limb movements performed while maintaining fixation produced as much interference with spatial working memory as reflexive saccades. These results suggest that the interference produced by eye movements is not the result of their visual consequences. Rather, all spatially directed movements appear to have similar effects on visuospatial working memory.     

What are human express saccades?

When a fixation point is removed 200 msec prior to target onset (the gap condition), human subjects are said to produce eye movements that have a short latency (80–120 msec), that form the early peak of a bimodal latency distribution, and that have been labeled “human express saccades” (see, e.g., Fischer, 1987; Fischer & Breitmeyer, 1987; Fischer & Ramsperger, 1984, 1986). In three experiments, we sought to obtain this express saccade diagnostic pattern in the gap condition, We orthogonally combined target location predictability with the presence versus absence of catch trials (Experiment 1). When target location was fixed and catch trials were not used, we found mostly anticipations. In the remaining conditions, where responses were under stimulus control, bimodality was not frequently observed, and, whether it was or not, latencies were not in the express saccade range. Using random target locations, we then varied stimulus luminance and the mode of stimulus presentation (LEDs vs. oscilloscope) in the gap and overlap (fixation is not removed) conditions (Experiment2). Bimodality was rarely observed, the gap effect (overlap minus gap reaction time) was additive with luminance, and only the brightest targets elicited saccades in the express range. When fixed locations and no catch trials were combined with latency feedback (Experiment 3), we observed many responses in the express saccade range and some evidence for bimodality, but the sudden introduction of catch trials revealed that many early responses were not under stimulus control. Humanscan make stimulus-controlled saccades that are initiated very rapidly (80–120 msec), but unless catch trials or choice reaction time is used, it is not possible to distinguish such saccades from anticipatory responses that are prepared in advance and timed to occur shortly after target onset. Because the express saccade diagnostic pattern is not a characteristic feature of human saccadic performance, we urge investigators to focus their attention on the robustgap effect     

Having to identify a target reduces latencies in prosaccades but not in antisaccades

In a princeps study, Trottier and Pratt (2005) showed that saccadic latencies were dramatically reduced when subjects were instructed to not simply look at a peripheral target (reflexive saccade) but to identify some of its properties. According to the authors, the shortening of saccadic reactions times may arise from a top-down disinhibition of the superior colliculus (SC), potentially mediated by the direct pathway connecting frontal/prefrontal cortex structures to the SC. Using a “cue paradigm” (a cue preceded the appearance of the target), the present study tests if the task instruction (Identify vs. Glance) also reduces the latencies of antisaccades (AS), which involve prefrontal structures. We show that instruction reduces latencies for prosaccade but not for AS. An AS requires two processes: the inhibition of a reflexive saccade and the generation of a voluntary saccade. To separate these processes and to better understand the task effect we also test the effect of the task instruction only on voluntary saccades. The effect still exists but it is much weaker than for reflexive saccades. The instruction effect closely depends on task demands in executive resources.     

Saccadic eye movements as indicators of cognitive function in older adults

Older adults appear to have greater difficulty ignoring distractions during day-to-day activities than younger adults. To assess these effects of age, the ability of adults aged between 50 and 80 years to ignore distracting stimuli was measured using the antisaccade and oculomotor capture tasks. In the antisaccade task, observers are instructed to look away from a visual cue, whereas in the oculomotor capture task, observers are instructed to look toward a colored singleton in the presence of a concurrent onset distractor. Index scores of the Repeatable Battery for the Assessment of Neuropsychological Status (RBANS) were compared with capture errors, and with prosaccade errors on the antisaccade task. A higher percentage of capture errors were made on the oculomotor capture tasks by the older members of the cohort compared to the younger members. There was a weak relationship between the attention index and capture errors, but the visuospatial/constructional index was the strongest predictor of prosaccade error rate in the antisaccade task. The saccade reaction times (SRTs) of correct initial saccades in the oculomotor capture task were poorly correlated with age, and with the neurospsychological tests, but prosaccade SRTs in both tasks moderately correlated with antisaccade error rate. These results were interpreted in terms of a competitive integration (or race) model. Any variable that reduces the strength of the top-down neural signal to produce a voluntary saccade, or that increases saccade speed, will enhance the likelihood that a reflexive saccade to a stimulus with an abrupt onset will occur.     

Initiation of bilateral responses in a kinesthetic reaction time task

Two experiments are presented in which different mechanisms controlling each limb in a bilateral arm response to a unilateral kinesthetic stimulus are postulated to occur. In Experiment 1, the limb serving as the stimulus are postulated to occur. In Experiment 1, the limb serving as the stimulus limb was described as operating with relative invariance by using a tightly coupled input-output reflexive pathway, whereas the nonstimulus limb appeared to be controlled by higher-order processing thought to be more susceptible to influences such as hemispheric specialization and stimulus expectancy. The differential control model was further tested in Experiment 2 by retaining the interhemispheric pathway of the unilateral kinesthetic stimulus but experimentally uncoupling the reflex mechanism from the stimulus side. Analyses of bilateral EMG premotor latencies under these conditions revealed that each response side can be controlled at separate levels-i.e., by a reflexive type mechanism or by higher-order processing when one of the response limbs is also the stimulus limb, both sides reflect behavior that is best described by an information processing type of voluntary control.     

Control of fixation duration in visual search and memory search: another look

In visual search a variable delay (up to 150 msec) between the beginning of each fixation and the onset of a search stimulus reduces the time (oculomotor latency) between stimulus onset and the subject's next saccadic eye movement. Two hypotheses for this effect of stimulus onset delay (SOD) were compared: first, process monitoring, that SOD simply serves as a warning interval to facilitate saccadic responses; and second, preprogramming, that saccades are preprogrammed at short SODs. In the first experiment SOD produced a decline in oculomotor latency in search similar to that seen in previous studies. In the second and third experiments, the size of the memory set in a Sternberg memory search paradigm was varied, or a mask flanking some of the search stimuli was used, to vary the processing time of each stimulus. Partial preprogramming of saccades at short delays would predict that increasing the processing time of individual stimuli would increase oculomotor latency at only short SODs. However, oculomotor latency increased equally at all SODs. In this search task, then, the SODs appeared to facilitate saccade initiation.     

Influence of endogenous and exogenous orientations of attention on inhibition of return in a cross-modal target-target aiming task

The authors conducted 2 experiments in which participants (N = 16 in each) executed successive unimanual aiming movements to target locations that were indicated by the onset of either an auditory or a visual stimulus. In Experiment 1 (exogenous orientation), inhibition of return (IOR) effects were observed, with reliable reaction time (RT) costs associated with movements returning to the same target and a trend toward larger IOR effects in left than in right space. There was no influence of stimulus modality on the magnitude of IOR. IOR was also observed in Experiment 2 (endogenous orientation), except the influence of stimulus modality reliably mediated those effect. In that case, IOR was evident only when the previous modality was visual and the current modality was auditory. Together, the results of those 2 experiments suggest that in situations in which 2 paired movements constitute the response criteria, IOR is both supramodal and lateralized to contralateral space.     

Initiation of Bilateral Responses in a Kinesthetic Reaction Time Task

Two experiments are presented in which different mechanisms controlling each limb in a bilateral arm response to a unilateral kinesthetic stimulus are postulated to occur. In Experiment 1, the limb serving as the stimulus limb was described as operating with relative invariance by using a tightly coupled input-output reflexive pathway, whereas the nonstimulus limb appeared to be controlled by higher-order processing thought to be more susceptible to influences such as hemispheric specialization and stimulus expectancy. The differential control model was further tested in Experiment 2 by retaining the intrahemispheric pathway of the unilateral kinesthetic stimulus but experimentally uncoupling the reflex mechanism from the stimulus side. Analyses of bilateral EMG premotor latencies under these conditions revealed that each response side can be controlled at separate levels —i.e., by a reflexive type mechanism or by higher-order processing when one of the response limbs is also the stimulus limb. When the stimulus is on the same side but not generated by either responding limb, both sides reflect behavior that is best described by an information processing type of voluntary control.     

Timing of reflexive visuospatial orienting in young, young-old, and old-old adults

This study examined adult age differences in reflexive orienting to two types of uninformative spatial cues: central arrows and peripheral onsets. In two experiments using a Posner cuing task, young adults (ages 18–28 years), young-old adults (60–74 years), and old-old adults (75–92 years) responded to targets that were preceded 100–1,000 ms earlier by a central arrow or a peripheral abrupt onset. In Experiment 1, the cue remained present upon target onset. Facilitation effects at short cue–target stimulus onset asynchronies (SOAs) were prolonged in duration for the two older groups relative to the young adults. At longer cue–target SOAs, inhibition of return (IOR) that was initiated by peripheral onset cues was observed in the performance of young adults but not in that of the two older groups. In Experiment 2, the cue was presented briefly and removed prior to target onset. The change in cue duration minimized age differences (particularly for young-old adults) in facilitation effects and led to IOR for all three age groups. The findings are consistent with the idea that attentional control settings change with age, with higher settings for older adults leading to delayed disengagement from spatial cues.     

Viewing behavior: Ocular and attentional disengagement

In five experiments, we examined the role of the ocular and attentional systems in determining saccadic latencies. Prior to making a saccade to a target stimulus, subjects were required to direct their attention to a foveal stimulus or to an eccentric stimulus. Either stimulus could be extinguished before the onset of the target. Saccadic latencies were shortest when the foveal stimulus was extinguished, regardless of whether it was attended or not. Control experiments showed that subjects were able to attend properly and that warning, arising from turning off a stimulus before target onset, could not completely account for the results. The results were discussed in terms of ocular disengagement, attentional disengagement, and joint ocular-attentional disengagement. It was concluded that an explanation emphasizing ocular disengagement provided the best account of the data.