Remembered Target Locations Research Articles

Neuronal activation sequences in lateral prefrontal cortex encode visuospatial working memory during virtual navigation

Working memory (WM) is the ability to maintain and manipulate information ‘in mind’. The neural codes underlying WM have been a matter of debate. We simultaneously recorded the activity of hundreds of neurons in the lateral prefrontal cortex of male macaque monkeys during a visuospatial WM task that required navigation in a virtual 3D environment. Here, we demonstrate distinct neuronal activation sequences (NASs) that encode remembered target locations in the virtual environment. This NAS code outperformed the persistent firing code for remembered locations during the virtual reality task, but not during a classical WM task using stationary stimuli and constraining eye movements. Finally, blocking NMDA receptors using low doses of ketamine deteriorated the NAS code and behavioral performance selectively during the WM task. These results reveal the versatility and adaptability of neural codes supporting working memory function in the primate lateral prefrontal cortex.

Egocentric cues influence the allocentric spatial memory of object configurations for memory-guided actions.

Allocentric and egocentric reference frames are used to code the spatial position of action targets in reference to objects in the environment, i.e., relative to landmarks (allocentric), or the observer (egocentric). Previous research investigated reference frames in isolation, for example, by shifting landmarks relative to the target and asking participants to reach to the remembered target location. Systematic reaching errors were found in the direction of the landmark shift and used as a proxy for allocentric spatial coding. Here, we examined the interaction of both allocentric and egocentric reference frames by shifting the landmarks as well as the observer. We asked participants to encode a three-dimensional configuration of balls and to reproduce this configuration from memory after a short delay followed by a landmark or an observer shift. We also manipulated the number of landmarks to test its effect on the use of allocentric and egocentric reference frames. We found that participants were less accurate when reproducing the configuration of balls after an observer shift, which was reflected in larger configurational errors. In addition, an increase in the number of landmarks led to a stronger reliance on allocentric cues and a weaker contribution of egocentric cues. In sum, our results highlight the important role of egocentric cues for allocentric spatial coding in the context of memory-guided actions.NEW & NOTEWORTHY Objects in our environment are coded relative to each other (allocentrically) and are thought to serve as independent and reliable cues (landmarks) in the context of unreliable egocentric signals. Contrary to this assumption, we demonstrate that egocentric cues alter the allocentric spatial memory, which could reflect recently discovered interactions between allocentric and egocentric neural processing pathways. Furthermore, additional landmarks lead to a higher contribution of allocentric and a lower contribution of egocentric cues.

Temporal dynamics of neural ensemble coding of remembered target location in the primate prefrontal cortex

Temporal dynamics of neural ensemble coding of remembered target location in the primate prefrontal cortex

Severe distortion in the representation of foveal visual image locations in short-term memory

The foveal visual image region provides the human visual system with the highest acuity. However, it is unclear whether such a high fidelity representational advantage is maintained when foveal image locations are committed to short-term memory. Here, we describe a paradoxically large distortion in foveal target location recall by humans. We briefly presented small, but high contrast, points of light at eccentricities ranging from 0.1 to 12°, while subjects maintained their line of sight on a stable target. After a brief memory period, the subjects indicated the remembered target locations via computer controlled cursors. The biggest localization errors, in terms of both directional deviations and amplitude percentage overshoots or undershoots, occurred for the most foveal targets, and such distortions were still present, albeit with qualitatively different patterns, when subjects shifted their gaze to indicate the remembered target locations. Foveal visual images are severely distorted in short-term memory.

Contributions of implicit and explicit memories to sensorimotor adaptation of movement extent during goal-directed reaching

We examined how implicit and explicit memories contribute to sensorimotor adaptation of movement extent during goal-directed reaching. Twenty subjects grasped the handle of a horizontal planar robot that rendered spring-like resistance to movement. Subjects made rapid “out-and-back” reaches to capture a remembered visual target at the point of maximal reach extent. The robot’s resistance changed unpredictably between reaches, inducing target capture errors that subjects attempted to correct from one trial to the next. Each subject performed over 400 goal-directed reaching trials. Some trials were performed without concurrent visual cursor feedback of hand motion. Some trials required self-assessment of performance between trials, whereby subjects reported peak reach extent on the most recent trial. This was done by either moving a cursor on a horizontal display (visual self-assessment), or by moving the robot’s handle back to the recalled location (proprioceptive self-assessment). Control condition trials performed either without or with concurrent visual cursor feedback of hand motion did not require self-assessments. We used step-wise linear regression analyses to quantify the extent to which prior reach errors and explicit memories of reach extent contribute to subsequent reach performance. Consistent with prior reports, providing concurrent visual feedback of hand motion increased reach accuracy and reduced the impact of past performance errors on future performance, relative to the corresponding no-vision control condition. By contrast, we found no impact of interposed self-assessment on subsequent reach performance or on how prior target capture errors influence subsequent reach performance. Self-assessments were biased toward the remembered target location and they spanned a compressed range of values relative to actual reach extents, demonstrating that declarative memories of reach performance systematically differed from actual performances. We found that multilinear regression could best account for observed data variability when the regression model included only implicit memories of prior reach performance; including explicit memories (self-assessments) in the model did not improve its predictive accuracy. We conclude therefore that explicit memories of prior reach performance do not contribute to implicit sensorimotor adaptation of movement extent during goal-directed reaching under conditions of environmental uncertainty.

Older Adults Show Reduced Spatial Precision but Preserved Strategy-Use During Spatial Navigation Involving Body-Based Cues.

Older adults typically perform worse on spatial navigation tasks, although whether this is due to degradation of memory or an impairment in using specific strategies has yet to be determined. An issue with some past studies is that older adults are tested on desktop-based virtual reality: a technology many report lacking familiarity with. Even when controlling for familiarity, these paradigms reduce the information-rich, three-dimensional experience of navigating to a simple two-dimensional task that utilizes a mouse and keyboard (or joystick) as means for ambulation. Here, we utilize a wireless head-mounted display and free ambulation to create a fully immersive virtual Morris water maze in which we compare the navigation of older and younger adults. Older and younger adults learned the locations of hidden targets from same and different start points. Across different conditions tested, older adults remembered target locations less precisely compared to younger adults. Importantly, however, they performed comparably from the same viewpoint as a switched viewpoint, suggesting that they could generalize their memory for the location of a hidden target given a new point of view. When we implicitly moved one of the distal cues to determine whether older adults used an allocentric (multiple landmarks) or beaconing (single landmark) strategy to remember the hidden target, both older and younger adults showed comparable degrees of reliance on allocentric and beacon cues. These findings support the hypothesis that while older adults have less precise spatial memories, they maintain the ability to utilize various strategies when navigating.

A neural network for online spike classification that improves decoding accuracy.

Separating neural signals from noise can improve brain-computer interface performance and stability. However, most algorithms for separating neural action potentials from noise are not suitable for use in real time and have shown mixed effects on decoding performance. With the goal of removing noise that impedes online decoding, we sought to automate the intuition of human spike-sorters to operate in real time with an easily tunable parameter governing the stringency with which spike waveforms are classified. We trained an artificial neural network with one hidden layer on neural waveforms that were hand-labeled as either spikes or noise. The network output was a likelihood metric for each waveform it classified, and we tuned the network's stringency by varying the minimum likelihood value for a waveform to be considered a spike. Using the network's labels to exclude noise waveforms, we decoded remembered target location during a memory-guided saccade task from electrode arrays implanted in prefrontal cortex of rhesus macaque monkeys. The network classified waveforms in real time, and its classifications were qualitatively similar to those of a human spike-sorter. Compared with decoding with threshold crossings, in most sessions we improved decoding performance by removing waveforms with low spike likelihood values. Furthermore, decoding with our network's classifications became more beneficial as time since array implantation increased. Our classifier serves as a feasible preprocessing step, with little risk of harm, that could be applied to both off-line neural data analyses and online decoding.NEW & NOTEWORTHY Although there are many spike-sorting methods that isolate well-defined single units, these methods typically involve human intervention and have inconsistent effects on decoding. We used human classified neural waveforms as training data to create an artificial neural network that could be tuned to separate spikes from noise that impaired decoding. We found that this network operated in real time and was suitable for both off-line data processing and online decoding.

Coding Locations Relative to One or Many Landmarks in Childhood.

Cognitive development studies how information processing in the brain changes over the course of development. A key part of this question is how information is represented and stored in memory. This study examined allocentric (world-based) spatial memory, an important cognitive tool for planning routes and interacting with the space around us. This is typically theorized to use multiple landmarks all at once whenever it operates. In contrast, here we show that allocentric spatial memory frequently operates over a limited spatial window, much less than the full proximal scene, for children between 3.5 and 8.5 years old. The use of multiple landmarks increases gradually with age. Participants were asked to point to a remembered target location after a change of view in immersive virtual reality. A k-fold cross-validation model-comparison selected a model where young children usually use the target location’s vector to the single nearest landmark and rarely take advantage of the vectors to other nearby landmarks. The comparison models, which attempt to explain the errors as generic forms of noise rather than encoding to a single spatial cue, did not capture the distribution of responses as well. Parameter fits of this new single- versus multi-cue model are also easily interpretable and related to other variables of interest in development (age, executive function). Based on this, we theorize that spatial memory in humans develops through three advancing levels (but not strict stages): most likely to encode locations egocentrically (relative to the self), then allocentrically (relative to the world) but using only one landmark, and finally, most likely to encode locations relative to multiple parts of the scene.

Impaired forward model updating in young adults with Tourette syndrome.

Current theories of motor control emphasize how the brain may use internal models of the body to ensure accurate planning and control of movements. One such internal model-a forward model-is thought to generate an estimate of the next motor state and/or the sensory consequences of an upcoming movement, thereby allowing movement errors to be monitored. In addition, forward models may provide a means by which to determine a sense of agency, i.e. the (conscious) sense of authorship and control over our actions. Tourette syndrome is a developmental neurological condition characterized by the occurrence of motor and phonic tics. The involuntary (or voluntary) nature of tics has been the subject of considerable debate, and it was recently argued that the presence of tics in Tourette syndrome could result in a blurring of any subjective boundary between voluntary and involuntary movements. In particular, it was proposed that the level of sensorimotor noise that accompanies tics may be particularly high in Tourette syndrome, and this may contribute to less efficient forward models used to determine agency. We investigated whether the internal monitoring of movements is impaired in individuals with Tourette syndrome, relative to a matched group of typically developing individuals, using a task that involved executing double-step aiming movements using a hand-held robot manipulandum. Participants were required on each trial to execute two movements in turn, each directed to a remembered target location without visual feedback. Importantly, we assumed that to perform accurately on the second (return) movement it would be necessary to update any forward model to take into account errors made during the first (outward) movement. Here we demonstrate that while the Tourette syndrome group were equally accurate, and no more variable, than the matched control group in executing aiming movements to the first (outward) target location, they were significantly less accurate, and exhibited greater movement variability, than controls when executing the second (return) movement. Furthermore, we show that for the return movement only, movement accuracy and movement variability were significantly predicted by the Tourette syndrome group's clinical severity scores. We interpret these findings as consistent with the view that individuals with Tourette syndrome may experience a reduction in the precision of the forward model estimates thought necessary for the accurate planning and control of movements.

Comparing Pupil Light Response Modulation between Saccade Planning and Working Memory.

The signature of spatial attention effects has been demonstrated through saccade planning and working memory. Although saccade planning and working memory have been commonly linked to attention, the comparison of effects resulting from saccade planning and working memory is less explored. It has recently been shown that spatial attention interacts with local luminance at the attended location. When bright and dark patch stimuli are presented simultaneously in the periphery, thereby producing no change in global luminance, pupil size is nonetheless smaller when the locus of attention overlaps with the bright, compared to the dark patch stimulus (referred to as the local luminance modulation). Here, we used the local luminance modulation to directly compare the effects of saccade planning and spatial working memory. Participants were required to make a saccade towards a visual target location (visual-delay) or a remembered target location (memory-delay) after a variable delay, and the bright and dark patch stimuli were presented during the delay period between target onset and go signal. Greater pupil constriction was observed when the bright patch, compared to the dark patch, was spatially aligned with the target location in both tasks. However, the effects were diminished when there was no contingency implemented between the patch and target locations, particularly in the memory-delay task. Together, our results suggest the involvement of similar, but not identical, attentional mechanisms through saccade planning and working memory, and highlight a promising potential of local pupil luminance responses for probing visuospatial processing.

Neural substrates for allocentric-to-egocentric conversion of remembered reach targets in humans.

Targets for goal-directed action can be encoded in allocentric coordinates (relative to another visual landmark), but it is not known how these are converted into egocentric commands for action. Here, we investigated this using a slow event-related fMRI paradigm, based on our previous behavioural finding that the allocentric-to-egocentric (Allo-Ego) conversion for reach is performed at the first possible opportunity. Participants were asked to remember (and eventually reach towards) the location of a briefly presented target relative to another visual landmark. After a first memory delay, participants were forewarned by a verbal instruction if the landmark would reappear at the same location (potentially allowing them to plan a reach following the auditory cue before the second delay), or at a different location where they had to wait for the final landmark to be presented before response, and then reach towards the remembered target location. As predicted, participants showed landmark-centred directional selectivity in occipital-temporal cortex during the first memory delay, and only developed egocentric directional selectivity in occipital-parietal cortex during the second delay for the 'Same cue' task, and during response for the 'Different cue' task. We then compared cortical activation between these two tasks at the times when the Allo-Ego conversion occurred, and found common activation in right precuneus, right presupplementary area and bilateral dorsal premotor cortex. These results confirm that the brain converts allocentric codes to egocentric plans at the first possible opportunity, and identify the four most likely candidate sites specific to the Allo-Ego transformation for reaches.

“Looking-at-nothing” during sequential sensorimotor actions: Long-term memory-based eye scanning of remembered target locations

Before acting humans saccade to a target object to extract relevant visual information. Even when acting on remembered objects, locations previously occupied by relevant objects are fixated during imagery and memory tasks – a phenomenon called “looking-at-nothing”. While looking-at-nothing was robustly found in tasks encouraging declarative memory built-up, results are mixed in the case of procedural sensorimotor tasks. Eye-guidance to manual targets in complete darkness was observed in a task practiced for days beforehand, while investigations using only a single session did not find fixations to remembered action targets. Here, it is asked whether looking-at-nothing can be found in a single sensorimotor session and thus independent from sleep consolidation, and how it progresses when visual information is repeatedly unavailable. Eye movements were investigated in a computerized version of the trail making test. Participants clicked on numbered circles in ascending sequence. Fifty trials were performed with the same spatial arrangement of 9 visual targets to enable long-term memory consolidation. During 50 consecutive trials, participants had to click the remembered target sequence on an empty screen. Participants scanned the visual targets and also the empty target locations sequentially with their eyes, however, the latter less precise than the former. Over the course of the memory trials, manual and oculomotor sequential target scanning became more similar to the visual trials. Results argue for robust looking-at-nothing during procedural sensorimotor tasks provided that long-term memory information is sufficient.

The Effects of Smoking Abstinence on Incentivized Spatial Working Memory

ABSTRACTBackground and Objective: Reward processing and working memory (WM) underlie value-based decision-making; consequently, joint examination of these systems may further our understanding of why smokers choose to smoke again following a quit attempt (relapse). While previous studies have demonstrated altered reward and WM function associated with nicotine exposure, little is known about the effects of abstinence on the joint function of these systems. The current study aims to address this gap. Method: Eighteen daily smokers were tested on a monetarily incentivized memory guided saccade (MGS) task on two separate, counterbalanced occasions, an abstinent and a non-abstinent session. The MGS task is a widely used metric of spatial working memory and enables precise quantification of the effects of rewards and nicotine exposure on behavior. Results: During the non-abstinent session, participants showed increased accuracy of the initial saccade towards the remembered target location on reward vs. neutral trials. Participants also showed increased accuracy of the final saccade towards the target, across incentive types, only during the non-abstinent condition. Discussion and Conclusions: Our observation that rewards improve the accuracy of the initial memory guided saccade during the non-abstinent but not abstinent condition extends a growing literature indicating reduced motivation towards monetary rewards during abstinence. Further, differences in the accuracy of the final corrective saccade during the non-abstinent but not the abstinent condition suggests smoking abstinence-related effects on WM precision beyond those related to incentive motivation (e.g., sustained attention). Significance: This work extends our fundamental understanding of smoking's effects on core affective and cognitive processes.

Causal inference in the updating and weighting of allocentric and egocentric information for spatial constancy during whole-body motion

It has been reported that the brain combines egocentric and allocentric information to update object positions after an intervening movement. Studies typically use discrete updating tasks (i.e., comparing pre- to post-movement target representations). Such approaches, however, cannot reveal how the brain would weigh the information in these reference frames during the intervening motion. A reasonable assumption is that objects with stable position over time would be more likely to be considered as a reliable allocentric landmark. But inferring whether an object is stable in space while the observer is moving involves attributing perceived changes in location to either the object's or the observer's displacement. Here, we tested this causal inference hypothesis by designing a continuous whole-body motion updating task. At the beginning of a trial, a target was presented for 500 ms, within a large visual frame. As soon as the target disappeared, participants were asked to move a cursor to this location by controlling a linear-guide mounted on the vestibular sled on which they were seated. Participants were translated sideways as soon as their reaching movement started, and they had to maintain the cursor on the remembered target location in space while being moved. During the sled motion, the frame would move with a velocity proportional to that of the sled (gain ranging from -0.7 to 0.7). Participants' responses showed a systematic bias in the direction of the frame displacement, one that increased with the difference between the frame and the sled velocity for small differences, but was decreasing for large differences. This bias pattern provides evidence for humans exploiting a dynamic Bayesian inference process with two causal structures to mediate the dynamic integration of allocentric and egocentric information in spatial updating. Meeting abstract presented at VSS 2017.

Let me be your guide: physical guidance improves spatial learning for older adults with simulated low vision.

Monitoring one's safety during low vision navigation demands limited attentional resources which may impair spatial learning of the environment. In studies of younger adults, we have shown that these mobility monitoring demands can be alleviated, and spatial learning subsequently improved, via the presence of a physical guide during navigation. The present study extends work with younger adults to an older adult sample with simulated low vision. We test the effect of physical guidance on improving spatial learning as well as general age-related changes in navigation ability. Participants walked with and without a physical guide on novel real-world paths in an indoor environment and pointed to remembered target locations. They completed concurrent measures of cognitive load on the trials. Results demonstrate an improvement in learning under low vision conditions with a guide compared to walking without a guide. However, our measure of cognitive load did not vary between guidance conditions. We also conducted a cross-age comparison and found support for age-related declines in spatial learning generally and greater effects of physical guidance with increasing age.

Multiple spatial representations interact to increase reach accuracy when coordinating a saccade with a reach.

Reaching is an essential behavior that allows primates to interact with the environment. Precise reaching to visual targets depends on our ability to localize and foveate the target. Despite this, how the saccade system contributes to improvements in reach accuracy remains poorly understood. To assess spatial contributions of eye movements to reach accuracy, we performed a series of behavioral psychophysics experiments in nonhuman primates (Macaca mulatta). We found that a coordinated saccade with a reach to a remembered target location increases reach accuracy without target foveation. The improvement in reach accuracy was similar to that obtained when the subject had visual information about the location of the current target in the visual periphery and executed the reach while maintaining central fixation. Moreover, we found that the increase in reach accuracy elicited by a coordinated movement involved a spatial coupling mechanism between the saccade and reach movements. We observed significant correlations between the saccade and reach errors for coordinated movements. In contrast, when the eye and arm movements were made to targets in different spatial locations, the magnitude of the error and the degree of correlation between the saccade and reach direction were determined by the spatial location of the eye and the hand targets. Hence, we propose that coordinated movements improve reach accuracy without target foveation due to spatial coupling between the reach and saccade systems. Spatial coupling could arise from a neural mechanism for coordinated visual behavior that involves interacting spatial representations.NEW & NOTEWORTHY How visual spatial representations guiding reach movements involve coordinated saccadic eye movements is unknown. Temporal coupling between the reach and saccade system during coordinated movements improves reach performance. However, the role of spatial coupling is unclear. Using behavioral psychophysics, we found that spatial coupling increases reach accuracy in addition to temporal coupling and visual acuity. These results suggest that a spatial mechanism to couple the reach and saccade systems increases the accuracy of coordinated movements.

Memory for Spatial Locations in a Patient with Near Space Neglect and Optic Ataxia: Involvement of the Occipitotemporal Stream.

Previous studies suggested that the occipitoparietal stream orients attention toward the near/lower space and is involved in immediate reaching, whereas the occipitotemporal stream orients attention toward the far/upper space and is involved in delayed reaching. In the present study, we investigated the role of the occipitotemporal stream in attention orienting and delayed reaching in a patient (GP) with bilateral damage to the occipitoparietal areas and optic ataxia. GP and healthy controls took part in three experiments. In the experiment 1, the participants bisected lines oriented along radial, vertical, and horizontal axes. GP bisected radial lines farther, and vertical lines more above, than the controls, consistent with an attentional bias toward the far/upper space and near/lower space neglect. The experiment 2 consisted of two tasks: (1) an immediate reaching task, in which GP reached target locations under visual control and (2) a delayed visual reaching task, in which GP and controls were asked to reach remembered target locations visually presented. We measured constant and variable distance and direction errors. In immediate reaching task, GP accurately reached target locations. In delayed reaching task, GP overshot remembered target locations, whereas the controls undershot them. Furthermore, variable errors were greater in GP than in the controls. In the experiment 3, GP and controls performed a delayed proprioceptive reaching task. Constant reaching errors did not differ between GP and the controls. However, variable direction errors were greater in GP than in the controls. We suggest that the occipitoparietal damage, and the relatively intact occipitotemporal region, produced in GP an attentional orienting bias toward the far/upper space (experiment 1). In turns, the attentional bias selectively shifted toward the far space remembered visual (experiment 2), but not proprioceptive (experiment 3), target locations. As a whole, these findings further support the hypothesis of an involvement of the occipitotemporal stream in delayed reaching. Furthermore, the observation that in both delayed reaching tasks the variable errors were greater in GP than in the controls suggested that in optic ataxia is present not only a visuo- but also a proprioceptivo-motor integration deficit.

Allocentric information is used for memory-guided reaching in depth: A virtual reality study

Previous research has demonstrated that humans use allocentric information when reaching to remembered visual targets, but most of the studies are limited to 2D space. Here, we study allocentric coding of memorized reach targets in 3D virtual reality. In particular, we investigated the use of allocentric information for memory-guided reaching in depth and the role of binocular and monocular (object size) depth cues for coding object locations in 3D space. To this end, we presented a scene with objects on a table which were located at different distances from the observer and served as reach targets or allocentric cues. After free visual exploration of this scene and a short delay the scene reappeared, but with one object missing (=reach target). In addition, the remaining objects were shifted horizontally or in depth. When objects were shifted in depth, we also independently manipulated object size by either magnifying or reducing their size. After the scene vanished, participants reached to the remembered target location on the blank table. Reaching endpoints deviated systematically in the direction of object shifts, similar to our previous results from 2D presentations. This deviation was stronger for object shifts in depth than in the horizontal plane and independent of observer-target-distance. Reaching endpoints systematically varied with changes in object size. Our results suggest that allocentric information is used for coding targets for memory-guided reaching in depth. Thereby, retinal disparity and vergence as well as object size provide important binocular and monocular depth cues.

The Effects of Restricted Peripheral Field-of-View on Spatial Learning while Navigating.

Recent work with simulated reductions in visual acuity and contrast sensitivity has found decrements in survey spatial learning as well as increased attentional demands when navigating, compared to performance with normal vision. Given these findings, and previous work showing that peripheral field loss has been associated with impaired mobility and spatial memory for room-sized spaces, we investigated the role of peripheral vision during navigation using a large-scale spatial learning paradigm. First, we aimed to establish the magnitude of spatial memory errors at different levels of field restriction. Second, we tested the hypothesis that navigation under these different levels of restriction would use additional attentional resources. Normally sighted participants walked on novel real-world paths wearing goggles that restricted the field-of-view (FOV) to severe (15°, 10°, 4°, or 0°) or mild angles (60°) and then pointed to remembered target locations using a verbal reporting measure. They completed a concurrent auditory reaction time task throughout each path to measure cognitive load. Only the most severe restrictions (4° and blindfolded) showed impairment in pointing error compared to the mild restriction (within-subjects). The 10° and 4° conditions also showed an increase in reaction time on the secondary attention task, suggesting that navigating with these extreme peripheral field restrictions demands the use of limited cognitive resources. This comparison of different levels of field restriction suggests that although peripheral field loss requires the actor to use more attentional resources while navigating starting at a less extreme level (10°), spatial memory is not negatively affected until the restriction is very severe (4°). These results have implications for understanding of the mechanisms underlying spatial learning during navigation and the approaches that may be taken to develop assistance for navigation with visual impairment.

No Evidence for a Saccadic Range Effect for Visually Guided and Memory-Guided Saccades in Simple Saccade-Targeting Tasks.

Saccades to single targets in peripheral vision are typically characterized by an undershoot bias. Putting this bias to a test, Kapoula [1] used a paradigm in which observers were presented with two different sets of target eccentricities that partially overlapped each other. Her data were suggestive of a saccadic range effect (SRE): There was a tendency for saccades to overshoot close targets and undershoot far targets in a block, suggesting that there was a response bias towards the center of eccentricities in a given block. Our Experiment 1 was a close replication of the original study by Kapoula [1]. In addition, we tested whether the SRE is sensitive to top-down requirements associated with the task, and we also varied the target presentation duration. In Experiments 1 and 2, we expected to replicate the SRE for a visual discrimination task. The simple visual saccade-targeting task in Experiment 3, entailing minimal top-down influence, was expected to elicit a weaker SRE. Voluntary saccades to remembered target locations in Experiment 3 were expected to elicit the strongest SRE. Contrary to these predictions, we did not observe a SRE in any of the tasks. Our findings complement the results reported by Gillen et al. [2] who failed to find the effect in a saccade-targeting task with a very brief target presentation. Together, these results suggest that unlike arm movements, saccadic eye movements are not biased towards making saccades of a constant, optimal amplitude for the task.