Neural correlates of inter- and intra-individual saccadic reaction time differences in the gap/overlap paradigm

To examine the neural correlates of contextually differing control mechanisms in saccade initiation, we studied 18 subjects who performed two saccade paradigms in a pseudo-random order, while their eye movements were recorded in the MRI scanner (1.5 T). In the gap task the fixation point was extinguished 200 ms before target onset, and in the overlap task the fixation point vanished 500 ms after target onset. Subjects were asked to maintain stable fixation in the fixation period and to quickly saccade to peripherally presented targets. Inter-individual activation differences were assessed using regression analyses at the second level, with mean saccadic reaction time (SRT) of subjects as a covariate. To identify brain regions varying with trial-by-trial changes in SRTs, we included SRTs as a parametric modulation regressor in the general linear model. All analyses were regions of interest based and were performed separately for the gap and overlap conditions. For the gap paradigm, we did not obtain activation in regions previously shown to be involved in preparatory processes with much longer gap periods. Interestingly, both inter- and intra-individual variability analyses revealed a positive correlation of activation in frontal and parietal eye-movement regions with SRTs, indicating that slower saccade performance is possibly associated with higher cortical control. For the overlap paradigm, the trial-by-trial variability analysis revealed a positive correlation of activation in the right opercular inferior frontal gyrus with SRTs, possibly linked to fixation-related processes that have to be overcome to perform a speeded saccade in presence of a fixation point.     

Express saccades in cat: effects of task and target modality

Saccadic eye movements to visual, auditory, and bimodal targets were measured in four adult cats. Bimodal targets were visual and auditory stimuli presented simultaneously at the same location. Three behavioral tasks were used: a fixation task and two saccadic tracking tasks (gap and overlap task). In the fixation task, a sensory stimulus was presented at a randomly selected location, and the saccade to fixate that stimulus was measured. In the gap and overlap tasks, a second target (hereafter called the saccade target) was presented after the cat had fixated the first target. In the gap task, the fixation target was switched off before the saccade target was turned on; in the overlap task, the saccade target was presented before the fixation target was switched off. All tasks required the cats to redirect their gaze toward the target (within a specified degree of accuracy) within 500 ms of target onset, and in all tasks target positions were varied randomly over five possible locations along the horizontal meridian within the cat's oculomotor range. In the gap task, a significantly greater proportion of saccadic reaction times (SRTs) were less than 125 ms, and mean SRTs were significantly shorter than in the fixation task. With visual targets, saccade latencies were significantly shorter in the gap task than in the overlap task, while, with bimodal targets, saccade latencies were similar in the gap and overlap tasks. On the fixation task, SRTs to auditory targets were longer than those to either visual or bimodal targets, but on the gap task, SRTs to auditory targets were shorter than those to visual or bimodal targets. Thus, SRTs reflected an interaction between target modality and task. Because target locations were unpredictable, these results demonstrate that cats, as well as primates, can produce very short latency goal-directed saccades.     

Relationship between directed visual attention and saccadic reaction times

Summary Saslow (1967) and Fischer and Ramsperger (1984) found that saccadic reaction time (SRT) depends on the interval between the fixation point offset and the target onset. Using a continuously visible fixation point, we asked whether a similar function would be obtained if subjects attended to a peripherally viewed point extinguished at variable intervals before or after the target onset. The interval was varied between -500ms (i.e., attention stimulus offset after saccade target onset = overlap trials) and 500ms (i.e., attention stimulus offset before saccade target onset = gap trials). The results show a constant mean SRT of about 240 ms for overlap trials, and a U-shaped function with a minimum of 140 ms, at a gap duration of 200 ms, for gap trials. These findings suggest that saccadic latencies do not depend on the cessation of fixation per se, but rather on the disengagement of attention from any location in the visual field. The time required for subjects to disengage their attention is approximately 100 ms. This disengaged state of attention — during which short latency (express) saccades can be made — can be sustained only for a gap duration of 300 ms. At longer gap durations mean SRTs increase again.     

Reaction times of vertical prosaccades and antisaccades in gap and overlap tasks

Horizontal saccadic reaction times (SRTs) have been extensively studied over the past 3 decades, concentrating on such topics as the gap effect, express saccades, training effects, and the role of fixation and attention. This study investigates some of these topics with regard to vertical saccades. The reaction times of vertical saccades of 13 subjects were measured using the gap and the overlap paradigms in the prosaccade task (saccade to the stimulus) and the antisaccade task (saccade in the direction opposite to the stimulus). In the gap paradigm, the initial fixation point (FP) was extinguished 200 ms before stimulus onset, while, in the overlap paradigm, the FP remained on during stimulus presentation. With the prosaccade overlap task, it was found that most subjects (10/13) — whether they were previously trained making horizontal saccades or naive — had significantly faster upward saccades compared with their downward saccades. One subject was faster in the downward direction and two were symmetrical. The introduction of the gap reduced the reaction times of the prosaccades, and express saccades were obtained in some naive and most trained subjects. This gap effect was larger for saccades made to the downward target. The strength of the updown asymmetry was more pronounced in the overlap as compared to the gap paradigm. With the antisaccade task, up-down asymmetries were much reduced. Express antisaccades were absent even with the gap paradigm, but reaction times were reduced as compared to the antisaccade overlap paradigm. There was a slight tendency for a larger gap effect of downward saccades. All subjects produced a certain number of erratic prosaccades in the antitaks, more with the gap than with the overlap paradigm. There was a significantly larger gap effect for the erratic prosaccades made to the downward, as compared to the upward, target, due to increased downward SRTs in the overlap paradigm. Three subjects trained in both the horizontal and the vertical direction showed faster SRTs and more express saccades in the horizontal directions as compared to the vertical. It is concluded that different parts of the visual field are differently organized with both directional and nondirectional components in saccade preparation.     

Auditory-visual interactions subserving goal-directed saccades in a complex scene

This study addresses the integration of auditory and visual stimuli subserving the generation of saccades in a complex scene. Previous studies have shown that saccadic reaction times (SRTs) to combined auditory-visual stimuli are reduced when compared with SRTs to either stimulus alone. However, these results have been typically obtained with high-intensity stimuli distributed over a limited number of positions in the horizontal plane. It is less clear how auditory-visual interactions influence saccades under more complex but arguably more natural conditions, when low-intensity stimuli are embedded in complex backgrounds and distributed throughout two-dimensional (2-D) space. To study this problem, human subjects made saccades to visual-only (V-saccades), auditory-only (A-saccades), or spatially coincident auditory-visual (AV-saccades) targets. In each trial, the low-intensity target was embedded within a complex auditory-visual background, and subjects were allowed over 3 s to search for and foveate the target at 1 of 24 possible locations within the 2-D oculomotor range. We varied systematically the onset times of the targets and the intensity of the auditory target relative to background [i.e., the signal-to-noise (S/N) ratio] to examine their effects on both SRT and saccadic accuracy. Subjects were often able to localize the target within one or two saccades, but in about 15% of the trials they generated scanning patterns that consisted of many saccades. The present study reports only the SRT and accuracy of the first saccade in each trial. In all subjects, A-saccades had shorter SRTs than V-saccades, but were more inaccurate than V-saccades when generated to auditory targets presented at low S/N ratios. AV-saccades were at least as accurate as V-saccades but were generated at SRTs typical of A-saccades. The properties of AV-saccades depended systematically on both stimulus timing and S/N ratio of the auditory target. Compared with unimodal A- and V-saccades, the improvements in SRT and accuracy of AV-saccades were greatest when the visual target was synchronous with or leading the auditory target, and when the S/N ratio of the auditory target was lowest. Further, the improvements in saccade accuracy were greater in elevation than in azimuth. A control experiment demonstrated that a portion of the improvements in SRT could be attributable to a warning-cue mechanism, but that the improvements in saccade accuracy depended on the spatial register of the stimuli. These results agree well with earlier electrophysiological results obtained from the midbrain superior colliculus (SC) of anesthetized preparations, and we argue that they demonstrate multisensory integration of auditory and visual signals in a complex, quasi-natural environment. A conceptual model incorporating the SC is presented to explain the observed data.     

Time-Window-of-Integration (TWIN) Model for Saccadic Reaction Time: Effect of Auditory Masker Level on Visual–Auditory Spatial Interaction in Elevation

Saccadic reaction time (SRT) to a visual target tends to be shorter when auditory stimuli are presented in close temporal and spatial proximity, even when subjects are instructed to ignore the auditory non-target (focused attention paradigm). Previous studies using pairs of visual and auditory stimuli differing in both azimuth and vertical position suggest that the amount of SRT facilitation decreases not with the physical but with the perceivable distance between visual target and auditory non-target. Steenken et al. (Brain Res 1220:150–156, 2008) presented an additional white-noise masker background of three seconds duration. Increasing the masker level had a diametrical effect on SRTs in spatially coincident versus disparate stimulus configurations: saccadic responses to coincident visual–auditory stimuli are slowed down, whereas saccadic responses to disparate stimuli are speeded up. Here we show that the time-window-of-integration model accounts for this observation by variation of a perceivable-distance parameter in the second stage of the model whose value does not depend on stimulus onset asynchrony between target and non-target.     

The influence of stimulus direction and eccentricity on pro- and anti-saccades in humans

We examined the sensory and motor influences of stimulus eccentricity and direction on saccadic reaction times (SRTs), direction-of-movement errors, and saccade amplitude for stimulus-driven (prosaccade) and volitional (antisaccade) oculomotor responses in humans. Stimuli were presented at five eccentricities, ranging from 0.5° to 8°, and in eight radial directions around a central fixation point. At 0.5° eccentricity, participants showed delayed SRT and increased direction-of-movement errors consistent with misidentification of the target and fixation points. For the remaining eccentricities, horizontal saccades had shorter mean SRT than vertical saccades. Stimuli in the upper visual field trigger overt shifts in gaze more easily and faster than in the lower visual field: prosaccades to the upper hemifield had shorter SRT than to the lower hemifield, and more anti-saccade direction-of-movement errors were made into the upper hemifield. With the exception of the 0.5° stimuli, SRT was independent of eccentricity. Saccade amplitude was dependent on target eccentricity for prosaccades, but not for antisaccades within the range we tested. Performance matched behavioral measures described previously for monkeys performing the same tasks, confirming that the monkey is a good model for the human oculomotor function. We conclude that an upper hemifield bias lead to a decrease in SRT and an increase in direction errors.     

Covert attention regulates saccadic reaction time by routing between different visual-oculomotor pathways

Covert attention modulates saccadic performance, e.g., the abrupt onset of a task-irrelevant visual stimulus grabs attention as measured by a decrease in saccadic reaction time (SRT). The attentional advantage bestowed by the task-irrelevant stimulus is short-lived: SRT is actually longer ~200 ms after the onset of a stimulus than it is when no stimulus appears, known as inhibition of return. The mechanism by which attention modulates saccadic reaction is not well-understood. Here, we propose two possible mechanisms: by selective routing of the visuomotor signal through different pathways (routing hypothesis) or by general modulation of the speed of visuomotor transformation (shifting hypothesis). To test them, we designed a cue gap paradigm in which a 100-ms gap was introduced between the fixation point disappearance and the target appearance to the conventional cued visual reaction time paradigm. The cue manipulated the location of covert attention, and the gap interval resulted in a bimodal distribution of SRT, with an early mode (express saccade) and a late mode (regular saccade). The routing hypothesis predicts changes in the proportion of express saccades vs. regular saccades, whereas the shifting hypothesis predicts a shift of SRT distribution. The addition of the cue had no effect on mean reaction time of express and regular saccades, but it changed the relative proportion of two modes. These results demonstrate that the covert attention modification of the mean SRT is largely attributed to selective routing between visuomotor pathways rather than general modulation of the speed of visuomotor transformation.     

Combined auditory and visual stimuli facilitate head saccades in the barn owl (Tyto alba)

The barn owl naturally responds to an auditory or visual stimulus in its environment with a quick head turn toward the source. We measured these head saccades evoked by auditory, visual, and simultaneous, co-localized audiovisual stimuli to quantify multisensory interactions in the barn owl. Stimulus levels ranged from near to well above saccadic threshold. In accordance with previous human psychophysical findings, the owl's saccade reaction times (SRTs) and errors to unisensory stimuli were inversely related to stimulus strength. Auditory saccades characteristically had shorter reaction times but were less accurate than visual saccades. Audiovisual trials, over a large range of tested stimulus combinations, had auditory-like SRTs and visual-like errors, suggesting that barn owls are able to use both auditory and visual cues to produce saccades with the shortest possible SRT and greatest accuracy. These results support a model of sensory integration in which the faster modality initiates the saccade and the slower modality remains available to refine saccade trajectory.     

Effects of pre-cues on voluntary and reflexive saccade generation I. Anti-cues for pro-saccades

Experiments on visual attention have employed both physical cues and verbal instructions to enable subjects to allocate attention at a location that becomes relevant within a perceptual or motor task some time later (cue lead time, CLT). In this study we have used valid visual peripheral cues (CLT between 100 and 700 ms) to indicate the direction and location of the next saccade. A cue is considered valid or invalid if its meaning with respect to the next saccade is correct or incorrect. A cue is called an anti- or pro-cue if the side of its presentation is opposite to or the same as the direction of the saccade required on a given trial. Correspondingly, a saccade is called an anti- or pro-saccade if it is directed to the side opposite to or the same as the stimulus presentation. A condition in which the cue and the stimulus are presented on opposite sides provides a simple way of dissociating voluntary attention allocation from automatic orienting. This paper considers the anti-cue pro-saccade task: the subjects were instructed to use the cue to direct attention to the opposite side, i.e. the location, where on valid trials the saccade target would occur. In the companion paper we have used the same physical condition, but we have reversed the instructions as to saccade direction and we have reversed the meaning of the cue, i.e. we designed a pro-cue anti-saccade task. In this first paper, the saccadic reaction times (SRTs) of pro-saccades of five adult subjects were measured in the gap paradigm (fixation point offset precedes target onset by 200 ms). With a CLT of 100 ms, valid anti-cues reduced the number of express saccades (i.e. saccades with SRTs in the range 80–120 ms) significantly compared with the control values (no cues). Valid anti-cues with increasingly long CLTs (100–700 ms) resulted in an increasing incidence of anticipatory saccades and saccades with longer SRTs (more than 120 ms), while the frequency of express saccades remained below the control value. When cue and saccade target were dissociated in location or in both location and direction, the effects of the cueing revealed a much lower spatial selectivity as compared to the effects that have been described for voluntary attention allocation by means of central cues. The results suggest that voluntary allocation of attention and cue-induced automatic orienting not only have different time courses but also have opposite effects on the generation of express saccades, and different spatial selectivities. A possible neuronal basis of these results is discussed considering related findings from electrophysiological studies in monkeys. Received: 27 March 1997 / Accepted: 17 December 1997     

Crossmodal integration in the primate superior colliculus underlying the preparation and initiation of saccadic eye movements

Saccades to combined audiovisual stimuli often have reduced saccadic reaction times (SRTs) compared with those to unimodal stimuli. Neurons in the intermediate/deep layers of the superior colliculus (dSC) are capable of integrating converging sensory inputs to influence the time to saccade initiation. To identify how neural processing in the dSC contributes to reducing SRTs to audiovisual stimuli, we recorded activity from dSC neurons while monkeys generated saccades to visual or audiovisual stimuli. To evoke crossmodal interactions of varying strength, we used auditory and visual stimuli of different intensities, presented either in spatial alignment or to opposite hemifields. Spatially aligned audiovisual stimuli evoked the shortest SRTs. In the case of low-intensity stimuli, the response to the auditory component of the aligned audiovisual target increased the activity preceding the response to the visual component, accelerating the onset of the visual response and facilitating the generation of shorter-latency saccades. In the case of high-intensity stimuli, the auditory and visual responses occurred much closer together in time and so there was little opportunity for the auditory stimulus to influence previsual activity. Instead, the reduction in SRT for high-intensity, aligned audiovisual stimuli was correlated with increased premotor activity (activity after visual burst but preceding saccade-aligned burst). These data provide a link between changes in neural activity related to stimulus modality with changes in behavior. They further demonstrate how crossmodal interactions are not limited to the initial sensory activity but can also influence premotor activity in the SC.     

Influence of stimulus eccentricity and direction on characteristics of pro- and antisaccades in non-human primates

The ability to inhibit reflexes in favor of goal-oriented behaviors is critical for optimal exploration and interaction with our environment. The antisaccade task can be used to investigate the ability of subjects to suppress a reflexive saccade (prosaccade) to a suddenly appearing visual stimulus and instead generate a voluntary saccade (antisaccade) to its mirror location. To understand the neural mechanisms required to perform this task, our lab has developed a non-human primate model. Two monkeys were trained on a task with randomly interleaved pro- and antisaccade trials, with the color of the central fixation point (FP) instructing the monkey to either make a prosaccade (red FP) or an antisaccade (green FP). In half of the trials, the FP disappeared 200 ms before stimulus presentation (gap condition) and in the remaining trials, the FP remained visible (overlap condition) during stimulus presentation. The effect of stimulus eccentricity and direction was examined by presenting the stimulus at one of eight different radial directions (0-360 degrees ) and five eccentricities (2, 4, 8, 10, and 16 degrees ). Antisaccades had longer saccadic reaction times (SRTs), more dysmetria, and lower peak velocities than prosaccades. Direction errors in the antisaccade task were more prevalent in the gap condition. The difference in mean SRT between correct pro- and antisaccades, the anti-effect, was greater in the overlap condition. The difference in mean SRT between the overlap and the gap condition, the gap effect, was larger for antisaccades than for prosaccades. The manipulation of stimulus eccentricity and direction influenced SRT and the proportion of direction errors. These results are comparable to human studies, supporting the use of this animal model for investigating the neural mechanisms subserving the generation of antisaccades.     

Event-related potentials and saccadic reaction times: effects of fixation point offset or change

Previous studies have shown that saccadic reaction times (SRTs) are reduced if the initial fixation point (FP) disappears 200 ms (gap period) before a peripheral target is presented. This gap saccade task is associated with a negative cortical potential at the end of the gap period. To determine whether the neural processes underlying this potential account for the reduction of SRTs during gap saccade tasks, we recorded event-related potentials (ERPs) in 19 subjects performing a gap saccade task (gap duration 200 ms), a warning saccade task (the color of the FP changed 200 ms prior to target appearance) and an overlap task (the FP remained visible during the trial). SRTs were shortest during the gap task, longest during the overlap task and intermediate during the warning task. The gap and warning tasks were accompanied by the same widespread negative cortical potential with a maximum at the time of stimulus presentation. These findings indicate that the warning effect mediated by the disappearance of the FP during gap saccade tasks is responsible for the gap negativity which was observed by several authors. Our findings of shorter SRTs during the gap task than the warning task, however, suggest that the gap has an additional effect that probably depends on subcortical mechanisms. Received: 01 June 1998 / Accepted: 12 March 1999     

Expression of a re-centering bias in saccade regulation by superior colliculus neurons

In previous studies of saccadic eye movement reaction time, the manipulation of initial eye position revealed a behavioral bias that facilitates the initiation of movements towards the central orbital position. An interesting hypothesis for this re-centering bias suggests that it reflects a visuo-motor optimizing strategy, rather than peripheral muscular constraints. Given that the range of positions that the eyes can take in the orbits delimits the extent of visual exploration by head-fixed subjects, keeping the eyes centered in the orbits may indeed permit flexible orienting responses to engaging stimuli. To investigate the influence of initial eye position on central processes such as saccade selection and initiation, we examined the activity of saccade-related neurons in the primate superior colliculus (SC). Using a simple reaction time paradigm wherein an initially fixated visual stimulus varying in position was extinguished 200 ms before the presentation of a saccadic target, we studied the relationship between initial eye position and neuronal activation in advance of saccade initiation. We found that the magnitude of the early activity of SC neurons, especially during the immediate pre-target period that followed the fixation stimulus disappearance, was correlated with changes in initial eye position. For the great majority of neurons, the pre-target activity increased with changes in initial eye position in the direction opposite to their movement fields, and it was also strongly correlated with the concomitant reduction in reaction time of centripetal saccades directed within their movement fields. Taking into account the correlation with saccadic reaction time, the relationship between neuronal activity and initial eye position remained significant. These results suggest that eye-position-dependent changes in the excitability of SC neurons could represent the neural substrate underlying a re-centering bias in saccade regulation. More generally, the low frequency SC pre-target activity could use eccentric eye position signals to regulate both when and which saccades are produced by promoting the emergence of a high frequency burst of activity that can act as a saccadic command. However, only saccades initiated within ~200 ms of target presentation were associated with SC pre-target activity. This eye-dependent pre-target activation mechanism therefore appears to be restricted to the initiation of saccades with relatively short reaction times, which specifically require the integrity of the SC. Electronic Publication     

Predictive remapping of visual features precedes saccadic eye movements

The frequent occurrence of saccadic eye movements raises the question of how information is combined across separate glances into a stable, continuous percept. Here I show that visual form processing is altered at both the current fixation position and the location of the saccadic target before the saccade. When human observers prepared to follow a displacement of the stimulus with the eyes, visual form adaptation was transferred from current fixation to the future gaze position. This transfer of adaptation also influenced the perception of test stimuli shown at an intermediate position between fixation and saccadic target. Additionally, I found a presaccadic transfer of adaptation when observers prepared to move their eyes toward a stationary adapting stimulus in peripheral vision. The remapping of visual processing, demonstrated here with form adaptation, may help to explain our impression of a smooth transition, with no temporal delay, of visual perception across glances.     

Age-related performance of human subjects on saccadic eye movement tasks

We measured saccadic eye movements in 168 normal human subjects, ranging in age from 5 to 79 years, to determine age-related changes in saccadic task performance. Subjects were instructed to look either toward (pro-saccade task) or away from (anti-saccade task) an eccentric target under different conditions of fixation. We quantified the percentage of direction errors, the time to onset of the eye movement (saccadic reaction time: SRT), and the metrics and dynamics of the movement itself (amplitude, peak velocity, duration) for subjects in different age groups. Young children (5–8 years of age) had slow SRTs, great intra-subject variance in SRT, and the most direction errors in the anti-saccade task. Young adults (20–30 years of age) typically had the fastest SRTs and lowest intra-subject variance in SRT. Elderly subjects (60–79 years of age) had slower SRTs and longer duration saccades than other subject groups. These results demonstrate very strong age-related effects in subject performance, which may reflect different stages of normal development and degeneration in the nervous system. We attribute the dramatic improvement in performance in the anti-saccade task that occurs between the ages of 5–15 years to delayed maturation of the frontal lobes. Received: 27 October 1997 / Accepted: 27 February 1998     

Evidence for interactions between target selection and visual fixation for saccade generation in humans

We examined the processes controlling selective orientation, specifically the processes required for generating saccadic eye movements in humans. Before a saccadic eye movement can be initiated, active visual fixation must be disengaged from the current point of fixation and a new target selected. We investigated whether these neural processes occur independently or interactively by devising a simple, multimodal choice reaction task in which subjects were asked to direct their gaze away from a central visual fixation target to an eccentric visual target while ignoring a simultaneous auditory distractor. Subjects had more difficulty suppressing incorrect movements toward the distractor when the fixation target was extinguished prior to onset of the eccentric target than when the fixation target remained illuminated during eccentric target presentation. Subjects with the shortest saccadic reaction times produced the most incorrect movements. These results support a recent hypothesis suggesting that the processes of disengaging active visual fixation and selecting a new saccade target are interrelated and arise, at least in part, from a change of activity within the superior colliculus.     

Behavioral properties of saccades generated as a choice response

The behavior characterizing choice response decision-making was studied in monkeys to provide background information for ongoing neurophysiological studies of the neural mechanisms underlying saccadic choice decisions. Animals were trained to associate a specific color from a set of colored visual stimuli with a specific spatial location. The visual stimuli (colored disks) appeared briefly at equal eccentricity from a central fixation position and then were masked by gray disks. The correct target association was subsequently cued by the appearance of a colored stimulus at the fixation point. The animal indicated its choice by saccading to the remembered location of the eccentric stimulus, which had matched the color of the cue. The number of alternative associations (NA) varied from 1 to 4 and remained fixed within a block of trials. After the training period, performance (percent correct responses) declined modestly as NA increased (on average 96, 93 or 84% correct for 1, 2 or 4 NA, respectively). Response latency increased logarithmically as a function of NA, thus obeying Hick’s law. The spatial extent of the learned association between color and location was investigated by rotating the array of colored stimuli that had remained fixed during the learning phase to various different angles. Error rates in choice saccades increased gradually as a function of the amount of rotation. The learned association biased the direction of the saccadic response toward the quadrant associated with the cue, but saccade direction was always toward one of the actual visual stimuli. This suggests that the learned associations between stimuli and responses were not spatially exact, but instead the association between color and location was distributed with declining strength from the trained locations. These results demonstrate that the saccade system in monkeys also displays the characteristic dependence on NA in choice response latencies, while more basic features of the eye movements are invariant from those in other tasks. The findings also provide behavioral evidence that spatially distributed regions are established for the sensory-to-motor associations during training which are later utilized for choice decisions.     

Time course of auditory masker effects: Tapping the locus of audiovisual integration?

In a focused attention paradigm, saccadic reaction time (SRT) to a visual target tends to be shorter when an auditory accessory stimulus is presented in close temporal and spatial proximity. Observed SRT reductions typically diminish as spatial disparity between the stimuli increases. Here a visual target LED (500 ms duration) was presented above or below the fixation point and a simultaneously presented auditory accessory (2 ms duration) could appear at the same or the opposite vertical position. SRT enhancement was about 35 ms in the coincident and 10 ms in the disparate condition. In order to further probe the audiovisual integration mechanism, in addition to the auditory non-target an auditory masker (200 ms duration) was presented before, simultaneous to, or after the accessory stimulus. In all interstimulus interval (ISI) conditions, SRT enhancement went down both in the coincident and disparate configuration, but this decrement was fairly stable across the ISI values. If multisensory integration solely relied on a feed-forward process, one would expect a monotonic decrease of the masker effect with increasing ISI in the backward masking condition. It is therefore conceivable that the relatively high-energetic masker causes a broad excitatory response of SC neurons. During this state, the spatial audio-visual information from multisensory association areas is fed back and merged with the spatially unspecific excitation pattern induced by the masker. Assuming that a certain threshold of activation has to be achieved in order to generate a saccade in the correct direction, the blurred joint output of noise and spatial audio-visual information needs more time to reach this threshold prolonging SRT to an audio-visual object.     

Startle effects on saccadic responses to emotional target stimuli

Startle stimuli elicit various physiological and cognitive responses. This study investigated whether acoustic startle stimuli affect saccadic reactions in an emotional pro‐ or antisaccade task. Startle probes were presented either 500 ms before or simultaneous with an imperative stimulus that indicated whether a saccade towards or away from positive, neutral, or negative peripheral target pictures had to be performed. Valence interacted with saccade direction according to an approach‐avoidance pattern of gaze behavior, with delayed prosaccades to negative targets and antisaccades away from positive targets. Acoustic startle stimuli preceding the presentation of peripheral target pictures speeded up the initiation saccades, irrespective of stimulus valence. Results indicate a speeding of cognitive‐motor processing by preceding startle stimuli.