Saccade dynamics during an online updating task change with healthy aging.

Goal-directed movements rely on the integration of both visual and motor information, especially during the online control of movement, to fluidly and flexibly control coordinated action. Eye-hand coordination typically plays an important role in goal-directed movements. As people age, various aspects of motor control and visual performance decline (Haegerstrom-Portnoy, Schneck, & Brabyn, 1999; Seidler et al., 2010), including an increase in saccade latencies (Munoz, Broughton, Goldring, & Armstrong, 1998). However, there is limited insight into how age-related changes in saccadic performance impact eye-hand coordination during online control. We investigated this question through the use of a target perturbation paradigm. Older and younger participants completed a perturbation task where target perturbations could occur either early (0 ms) or later (200 ms) after reach onset. We analyzed reach correction latencies and the frequency of the reach correction, coupled with analyses of saccades across all stages of movement. Older participants had slower correction latencies and initiated corrections less frequently compared to younger participants, with this trend being exacerbated in the later (200 ms) target perturbation condition. Older participants also produced slower saccade latencies toward both the initial target and the perturbed target. For trials in which a correction occurred to a late perturbation, touch responses were more accurate when there was more time between the saccade landing and the touch. Altogether, our results suggest that these age-related effects may be due to the delayed acquisition of visual and oculomotor information used to inform the reaching movement, stemming from the increase in saccade latencies before and after target perturbation.

A Depth-Dependent Integrated VF Simulation for Analysis and Visualization of Glaucomatous VF Defects.

Purpose: Visual fields (VF) are measured monocularly at a single depth, yet real-life activities require people to interact with objects binocularly at multiple depths. To better characterize visual functioning in clinical vision conditions such as glaucoma, analyzing visual impairment in a depth-dependent fashion is required. We developed a depth-dependent integrated VF (DD-IVF) simulation and demonstrated its usefulness by evaluating DD-IVF defects associated with 12 glaucomatous archetypes of 24-2 VF. Methods: The 12 archetypes included typical variants of superior and inferior nasal steps, arcuate and altitudinal defects, temporal wedge, biarcuate, and intact VFs. DD-IVF simulation maps the monocular 24-2 VF archetypes to binocular ones as a function of depth by incorporating three parameters of fixation, object, and interpupillary distances. At each location and depth plane, sensitivities are linearly interpolated from corresponding locations in monocular VF and returned as the higher value of the two. Results: The simulation produced 144 DD-IVFs for multiple depths from combinations of 12 glaucomatous archetypes. The DD-IVFs are included as a Shiny app in the binovisualfields package. The number of impaired locations in the DD-IVFs varied according to the overlap of VF loss between eyes. Conclusions: Our DD-IVF program revealed binocular functional visual defects associated with glaucomatous archetypes of the 24-2 pattern and is designed to do the same for empirically measured VFs. The comparison of identified visual impairments across depths may be informative for future empirical exploration of functional visual impairments in depth in glaucoma and other conditions leading to bilateral VF loss. Translational Relevance: Our DD-IVF program can reveal depth-dependent functional visual defects for clinical vision conditions where 24-2 test patterns are available.

The effect of age and perturbation time on online control during rapid pointing.

Visual and proprioceptive information is used differently at different phases of a reach. The time at which a target perturbation occurs during a reach therefore has a significant impact on how an individual can compensate for this perturbation though online control. With healthy ageing, there are notable changes to both sensory and motor control that impact motor performance. However, how the online control process changes with age is not yet fully understood. We used a target perturbation paradigm and manipulated the time at which a target perturbation occurred during the reach to investigate how healthy ageing impacts sensorimotor control. We measured how the latency of the correction and the magnitude of the corrective response changed with perturbation time and quantified the difference across groups using a percentage difference measure. For both groups, online corrections to early perturbations were more easily accounted for than those to late perturbations, despite late perturbations eliciting faster correction latencies. While there was no group difference in accuracy, older participants were slower overall and produced a correction to a change in target location proportionally less often despite similar correction latencies. We speculate that the differences in the time during the reach that the correction is first identified may explain the differences in correction latencies observed between the perturbation time conditions.

The spatial and temporal properties of attentional selectivity for saccades and reaches.

The preparation and execution of saccades and goal-directed movements elicits an accompanying shift in attention at the locus of the impending movement. However, some key aspects of the spatiotemporal profile of this attentional shift between eye and hand movements are not resolved. While there is evidence that attention is improved at the target location when making a reach, it is not clear how attention shifts over space and time around the movement target as a saccade and a reach are made to that target. Determining this spread of attention is an important aspect in understanding how attentional resources are used in relation to movement planning and guidance in real world tasks. We compared performance on a perceptual discrimination paradigm during a saccade-alone task, reach-alone task, and a saccade-plus-reach task to map the temporal profile of the premotor attentional shift at the goal of the movement and at three surrounding locations. We measured performance relative to a valid baseline level to determine whether motor planning induces additional attentional facilitation compared to mere covert attention. Sensitivity increased relative to movement onset at the target and at the surrounding locations, for both the saccade-alone and saccade-plus-reach conditions. The results suggest that the temporal profile of the attentional shift is similar for the two tasks involving saccades (saccade-alone and saccade-plus-reach tasks), but is very different when the influence of the saccade is removed. In this case, performance in the saccade-plus-reach task reflects the lower sensitivity observed when a reach-alone task is being conducted. In addition, the spatial profile of this spread of attention is not symmetrical around the target. This suggests that when a saccade and reach are being planned together, the saccade drives the attentional shift, and the reach-alone carries little attentional weight.

Changes to online control and eye-hand coordination with healthy ageing.

Goal directed movements are typically accompanied by a saccade to the target location. Online control plays an important part in correction of a reach, especially if the target or goal of the reach moves during the reach. While there are notable changes to visual processing and motor control with healthy ageing, there is limited evidence about how eye-hand coordination during online updating changes with healthy ageing. We sought to quantify differences between older and younger people for eye-hand coordination during online updating. Participants completed a double step reaching task implemented under time pressure. The target perturbation could occur 200, 400 and 600 ms into a reach. We measured eye position and hand position throughout the trials to investigate changes to saccade latency, movement latency, movement time, reach characteristics and eye-hand latency and accuracy. Both groups were able to update their reach in response to a target perturbation that occurred at 200 or 400 ms into the reach. All participants demonstrated incomplete online updating for the 600 ms perturbation time. Saccade latencies, measured from the first target presentation, were generally longer for older participants. Older participants had significantly increased movement times but there was no significant difference between groups for touch accuracy. We speculate that the longer movement times enable the use of new visual information about the target location for online updating towards the end of the movement. Interestingly, older participants also produced a greater proportion of secondary saccades within the target perturbation condition and had generally shorter eye-hand latencies. This is perhaps a compensatory mechanism as there was no significant group effect on final saccade accuracy. Overall, the pattern of results suggests that online control of movements may be qualitatively different in older participants.

The Effect of Locomotion on Early Visual Contrast Processing in Humans.

Most of our knowledge about vision comes from experiments in which stimuli are presented to immobile human subjects or animals. In the case of human subjects, movement during psychophysical, electrophysiological, or neuroimaging experiments is considered to be a source of noise to be eliminated. Animals used in visual neuroscience experiments are typically restrained and, in many cases, anesthetized. In reality, however, vision is often used to guide the motion of awake, ambulating organisms. Recent work in mice has shown that locomotion elevates visual neuronal response amplitudes (Niell and Stryker, 2010; Erisken et al., 2014; Fu et al., 2014; Lee et al., 2014; Mineault et al., 2016) and reduces long-range gain control (Ayaz et al., 2013). Here, we used both psychophysics and steady-state electrophysiology to investigate whether similar effects of locomotion on early visual processing can be measured in humans. Our psychophysical results show that brisk walking has little effect on subjects' ability to detect briefly presented contrast changes and that co-oriented flankers are, if anything, more effective masks when subjects are walking. Our electrophysiological data were consistent with the psychophysics indicating no increase in stimulus-driven neuronal responses while walking and no reduction in surround suppression. In summary, we have found evidence that early contrast processing is altered by locomotion in humans but in a manner that differs from that reported in mice. The effects of locomotion on very low-level visual processing may differ on a species-by-species basis and may reflect important differences in the levels of arousal associated with locomotion.SIGNIFICANCE STATEMENT Mice are the current model of choice for studying low-level visual processing. Recent studies have shown that mouse visual cortex is modulated by behavioral state: primary visual cortex neurons in locomoting mice tend to be more sensitive and less influenced by long-range gain control. Here, we tested these effects in humans by measuring psychophysical detection thresholds and electroencephalography (EEG) responses while subjects walked on a treadmill. We found no evidence of increased contrast sensitivity or reduced surround suppression in walking humans. Our data show that fundamental measurements of early visual processing differ between humans and mice and this has important implications for recent work on the links among arousal, behavior, and vision in these two species.

The profile of attention differs between locations orthogonal to and in line with reach direction.

People make movements in a variety of directions when interacting with the world around them. It has been well documented that attention shifts to the goal of an upcoming movement, whether the movement is a saccade or a reach. However, recent evidence suggests that the direction of a movement may influence the spatial spread of attention (Stewart & Ma-Wyatt, 2015, Journal of Vision, 15(5), 10). We investigated whether the spatiotemporal profile of attention differs depending on where that location is situated relative to the direction of movement, and if this pattern is consistent across different movement effectors. We compared attentional facilitation at locations in line with or orthogonal to the movement, for reach-only, reach-plus-saccade, and saccade-only conditions. Results show that the spatiotemporal profile of attention differs across different movement combinations, and is also different at target locations orthogonal to and in line with the movement direction. Specifically, when a reach alone was made, there was a general decrease in attention at all locations during the movement and a general increase in attention at all locations with a saccade only. However, the concurrent reach and saccade condition showed a premovement attentional facilitation at locations orthogonal to movement direction, but not those in line with the movement direction. These results suggest attentional guidance may be more important at differing time points, depending on the type of movement.

The structure of sequential effects.

There is a long history of research into sequential effects, extending more than one hundred years. The pattern of sequential effects varies widely with both experimental conditions as well as for different individuals performing the same experiment. Yet this great diversity of results is poorly understood, particularly with respect to individual variation, which save for some passing mentions has largely gone unreported in the literature. Here we seek to understand the way in which sequential effects vary by identifying the causes underlying the differences observed in sequential effects. In order to achieve this goal we perform principal component analysis on a dataset of 158 individual results from participants performing different experiments with the aim of identifying hidden variables responsible for sequential effects. We find a latent structure consisting of 3 components related to sequential effects-2 main and 1 minor. A relationship between the 2 main components and the separate processing of stimuli and of responses is proposed on the basis of previous empirical evidence. It is further speculated that the minor component of sequential effects arises as the consequence of processing delays. Independently of the explanation for the latent variables encountered, this work provides a unified descriptive model for a wide range of different types of sequential effects previously identified in the literature. In addition to explaining individual differences themselves, it is demonstrated how the latent structure uncovered here is useful in understanding the classical problem of the dependence of sequential effects on the interval between successive stimuli.

The spatiotemporal characteristics of the attentional shift relative to a reach.

While the attentional shift preceding a saccadic eye movement has been well documented, the mechanisms surrounding the attentional shift preceding a reach are not well understood. It is unknown whether these mechanisms may be the same as those used in perceptual tasks, or those used in the planning of a saccade. We mapped the spatiotemporal properties of attention relative to a reach to determine the time course of attentional facilitation for hand movements alone. Participants had to reach toward a target and during the reach a perceptual probe could appear at one of six locations around the target, and at nine temporal offsets relative to the cue. Results showed a consistent pattern of facilitation in the planning stages of the reach, with attention increasing and then reaching a plateau during the completion of the movement before dropping off. These results demonstrate that planning a hand movement necessitates a shift in attention across the visual field around 150 ms before the onset of a reach. While these results are broadly consistent with the results of experiments mapping attentional shifts for saccades, the spatiotemporal profile of facilitation found shows that reaching without a concurrent eye movement also causes shifts in attention across the visual field. These results also suggest that the profile of the attentional shift preceding and during a hand movement is different at different locations across the visual field.

The effects of ageing and visual field loss on pointing to visual targets.

PURPOSE: To investigate the effect of ageing on visuomotor function and subsequently evaluate the effect of visual field loss on such function in older adults. METHODS: Two experiments were performed: 1) to determine the effect of ageing on visual localisation and subsequent pointing precision, and 2) to determine the effect of visual field loss on these outcome measures. For Experiment 1, we measured visual localisation and pointing precision radially at visual eccentricities of 5, 10 and 15° in 25 older (60-72 years) and 25 younger (20-31 years) adults. In the pointing task, participants were asked to point to a target on a touchscreen at a natural pace that prioritised accuracy of the touch. In Experiment 2, a subset of these tasks were performed at 15° eccentricity under both monocular and binocular conditions, by 8 glaucoma (55-76 years) and 10 approximately age-matched controls (61-72 years). RESULTS: Visual localisation and pointing precision was unaffected by ageing (p>0.05) and visual field loss (p>0.05), although movement time was increased in glaucoma (p = 0.01). CONCLUSION: Visual localisation and pointing precision to high contrast stimuli within the central 15° of vision are unaffected by ageing. Even in the presence of significant visual field loss, older adults with glaucoma are able perform such tasks with reasonable precision provided the target can be perceived and movement time is not restricted.

The distribution of spatial attention changes with task demands during goal-directed reaching.

Goal-directed movements are commonly used to allow humans to interact with their environment. When making a goal-directed movement in a natural environment, there are many competing stimuli. It is therefore important to understand how making a goal-directed movement could be impacted by the need to divide attention between the movement and competing stimuli. We used a dual-task paradigm to investigate the sharing of attentional resources between a search task in central vision and a peripheral pointing task completed concurrently. Results suggest some degree of shared attentional resources between these two tasks with performance on both central and peripheral tasks degraded under dual-task conditions. Movement latency, but not movement time, was also affected by dual-task conditions. Altogether, the results suggest that there is a cost to reach performance if attention is engaged away from the movement goal. Interestingly, this cost is associated with movement planning rather than execution.

Delays in using chromatic and luminance information to correct rapid reaches.

People can use feedback to make online corrections to movements but only if there is sufficient time to integrate the new information and make the correction. A key variable in this process is therefore the speed at which the new information about the target location is coded. Conduction velocities for chromatic signals are lower than for achromatic signals so it may take longer to correct reaches to chromatic stimuli. In addition to this delay, the sensorimotor system may prefer achromatic information over the chromatic information as delayed information may be less valuable when movements are made under time pressure. A down-weighting of chromatic information may result in additional latencies for chromatically directed reaches. In our study, participants made online corrections to reaches to achromatic, (L-M)-cone, and S-cone stimuli. Our chromatic stimuli were carefully adjusted to minimize stimulation of achromatic pathways, and we equated stimuli both in terms of detection thresholds and also by their estimated neural responses. Similar stimuli were used throughout the subjective adjustments and final reaching experiment. Using this paradigm, we found that responses to achromatic stimuli were only slightly faster than responses to (L-M)-cone and S-cone stimuli. We conclude that the sensorimotor system treats chromatic and achromatic information similarly and that the delayed chromatic responses primarily reflect early conduction delays.

Eye-hand coordination while pointing rapidly under risk.

Humans make rapid, goal-directed movements to interact with their environment. Saccadic eye movements usually accompany rapid hand movements, suggesting neural coupling, although it remains unclear what determines the strength of the coupling. Here, we present evidence that humans can alter eye-hand coordination in response to risk associated with endpoint variability. We used a paradigm in which human participants were forced to point rapidly under risk and were penalized or rewarded depending on the hand movement outcome. A separate reward schedule was employed for relative saccadic endpoint position. Participants received a monetary reward proportional to points won. We present a model that defines optimality of eye-hand coordination for this task depending on where the hand lands relative to the eye. A comparison of the results and model predictions showed that participants could optimize performance to maximize gain in some conditions, but not others. Participants produced near-optimal results when no feedback was given about relative saccade location and when negative feedback was provided for large distances between the saccade and hand. Participants were sub-optimal when given negative feedback for saccades very close to the hand endpoint. Our results suggest that eye-hand coordination is flexible when pointing rapidly under risk, but final eye position remains correlated with finger location.

The wallpaper illusion explained.

In the wallpaper illusion, a repetitive pattern appears to shift from one depth plane to the plane nearest fixation. We measured the timing of this shift for a 6 degrees wide, 3-cpd sinusoidal grating presented in a rectangular envelope; the edges (envelope) of the grating were presented at 20 arcmin of disparity (one period) behind the fixation plane. We asked observers to signal when the segment appeared to move from the edge plane forward to the fixation plane. Initially, the shift from the edge plane took 4-6 s, but after many trials, the shift became faster. Additional experiments demonstrated that the envelope was adapting, thereby permitting the alternative match. Our measurements for a range of spatial frequencies and disparities showed that these shifts to the fixation plane occurred only if the envelope disparity was more than one-half period of the carrier; that is, phase disparity >180 degrees. We also found that stereoacuity for the initial envelope-based match was poor, as might be expected for a target presented far off the fixation plane. However, once the perceived shift in depth occurred, stereoacuity improved fivefold without any change in the physical stimulus. We speculate that access to the most sensitive V1 neurons depends on the extrastriate processes that determine perceived depth--in this case, second-order envelope mechanisms.

Visual information throughout a reach determines endpoint precision.

People make rapid, goal-directed movements to interact with their environment. Because these movements have consequences, it is important to be able to control them with a high level of precision and accuracy. Our hypothesis is that vision guides rapid hand movements, thereby enhancing their accuracy and precision. To test this idea, we asked observers to point to a briefly presented target (110 ms). We measured the impact of visual information on endpoint precision by using a shutter to close off view of the hand 50, 110 and 250 ms into the reach. We found that precision was degraded if the view of the hand was restricted at any time during the reach, despite the fact that the target disappeared long before the reach was completed. We therefore conclude that vision keeps the hand on the planned trajectory. We then investigated the effects of a perturbation of target position during the reach. For these experiments, the target remained visible until the reach was completed. The target position was shifted at 110, 180 or 250 ms into the reach. Early shifts in target position were easily compensated for, but late shifts led to a shift in the mean position of the endpoints; observers pointed to the center of the two locations, as a kind of best bet on the position of the target. Visual information is used to guide the hand throughout a reach and has a significant impact on endpoint precision.

Initial visual information determines endpoint precision for rapid pointing.

We investigated how visual noise in the initial estimate of target location affects precision for rapid pointing. Visual localization thresholds (an error measure) rise systematically with eccentricity, doubling at eccentricities of a degree or less. Previous work, which we confirmed, has shown that the precision of pointing, measured by the standard deviation, to a single isolated target is relatively constant over small lateral extents near the midline, and that pointing error is substantially larger than visual error. We used target uncertainty (randomly chosen locations) to greatly increase visual noise so that we could explore the influence of visual noise on pointing error. We compared precision for comparable visual and pointing tasks as a function of target eccentricity. The target was presented for 110 ms at one of eight isoeccentric locations, chosen at random. Under these conditions, pointing error increased significantly with increasing target eccentricity. Beyond 4 degrees eccentricity, visual thresholds and pointing error were identical. Even when the target remained visible until the movement was completed, initial target eccentricity affected pointing error. The quality of visual information varies with task demands, and therefore so does its influence on endpoint precision. Our results demonstrate that the initial visual information about target location can limit endpoint precision, even over as small a range as 12 degrees in the central visual field (a lateral extent of +/-8.5 cm at the midline).

Seeing and ballistic pointing at perisaccadic targets.

We studied the effects of visual references and the level of illumination on the localization of stimuli flashed briefly near the start of saccades. A translucent shutter made it possible to remove visual references, but admit light, at different times after saccadic onset. The results show that post-saccadic visual references are not necessary for compression: a consistent compression of verbally reported relative stimulus distances is found at all shutter latencies and at all post-shutter levels of illumination. They also show that positions indicated by blind pointing show no compression except when visual references remain in view for a substantial time after saccades. These results confirm that the visual system uses multiple representations of space and suggest that it weights them differently for different tasks and different viewing conditions. No single map is used exclusively for conscious perception or for motor action, and conscious perception is always subject to compression at the time of saccades.

Contrast configuration influences grouping in apparent motion.

We investigated whether the same principles that influence grouping in static displays also influence grouping in apparent motion. Using the Ternus display, we found that the proportion of group motion reports was influenced by changes in contrast configuration. Subjects made judgments of completion of these same configurations in a static display. Generally, contrast configurations that induced a high proportion of group motion responses were judged as more 'complete' in static displays. Using a stereo display, we then tested whether stereo information and T-junction information were critical for this increase in group motion. Perceived grouping was consistently higher for same contrast polarity configurations than for opposite contrast polarity configurations, regardless of the presence of stereo information or explicit T-junctions. Thus, while grouping in static and moving displays showed a similar dependence on contrast configuration, motion grouping showed little dependence on stereo or T-junction information.

Saccades actively maintain perceptual continuity.

People make saccades--rapid eye movements to a new fixation--approximately three times per second. This would seemingly disrupt perceptual continuity, yet our brains construct a coherent, stable view of the world from these successive fixations. There is conflicting evidence regarding the effects of saccades on perceptual continuity: some studies report that they are disruptive, with little information carryover between saccades; others report that carryover is substantial. Here we show that saccades actively contribute to perceptual continuity in humans in two different ways. When bistable stimuli are presented intermittently, saccades executed during the blank interval shorten the duration of states of ambiguous figures, indicating that saccades can erase immediately past perceptual states. On the other hand, they prolong the McCollough effect, indicating that saccades strengthen learned contingencies. Our results indicate that saccades help, rather than hinder, perceptual continuity.

Developmental dyslexia in different languages: language-specific or universal?

Most of the research on developmental dyslexia comes from English-speaking countries. However, there is accumulating evidence that learning to read English is harder than learning to read other European orthographies (Seymour, Aro, & Erskine, 2003). These findings therefore suggest the need to determine whether the main English findings concerning dyslexia can be generalized to other European orthographies, all of which have less irregular spelling-to-sound correspondences than English. To do this, we conducted a study with German- and English-speaking children (n=149) in which we investigated a number of theoretically important marker effects of the reading process. The results clearly show that the similarities between dyslexic readers using different orthographies are far bigger than their differences. That is, dyslexics in both countries exhibit a reading speed deficit, a nonword reading deficit that is greater than their word reading deficit, and an extremely slow and serial phonological decoding mechanism. These problems were of similar size across orthographies and persisted even with respect to younger readers that were at the same reading level. Both groups showed that they could process larger orthographic units. However, the use of this information to supplement grapheme-phoneme decoding was not fully efficient for the English dyslexics.