"Warm," "cool," and the colors.

Participants judged affective cooler/warmer gradients around a 12-step color circle. Each pair of adjacent colors was presented twice (left-right reversed), all in random order. Participants readily performed the task, but their settings do not correlate very well. Individual responses were compared with a small number of canonical templates. For a little less than one-half of the participants responses or judgements correlate with such a template. We find a warm pole (in the orange environment) and a cool pole (in the teal environment) connected with two tracks that tend to have one or more gaps or weak, even inverted links. We conclude that the common artistic cool-warm polarity is only weakly reflected in responses of our observers. If it does, the observers apparently use categorical warm and cool poles and may be uncertain in relating adjacent hue steps along the 12-step color circle.

Context effects on the perception of saturation of fruit colors in still-life paintings.

Still-life painters, especially of the so-called Golden Age (17th century) in the Netherlands, are famous for their masterful techniques of rendering reality. Their amazing abilities to depict different material properties of fruits and flowers are stunning. But how important are these careful arrangements of different objects for the perception of an individual item? Is the perceived color saturation of a single fruit influenced by its surrounding context? We selected fruits in still-life paintings as stimuli to investigate whether and how perceived saturations of fruits were affected by their original contexts. In our study, we focused especially on effects of five context properties: complementary colors, chromatic and luminance contrast, object overlap, and surround variance. Six fruit varieties depicted in high-quality digital reproductions of 48 classic and eight varieties in 64 more recent, modern still-life paintings were selected. In a single trial, eight images of fruits of the same variety appeared on a neutral gray background; half were single fruit cutouts, and the other half were the same fruits embedded in their circular contexts. Fifteen participants ranked all eight images according to perceived color saturations of the fruits. Saturation ratings showed a high agreement of 77%. Surrounding contexts led to an increase in perceived saturation of central fruits. This effect was mainly driven by object overlap, the presence of the central fruit type also in the context, and surround variance. Chroma contrast between fruits and contexts decreased saturation significantly. No significant context effects were found for complementary colors or luminance contrast. Our results show that in paintings, many of the cues that are usually experimentally isolated occur in interesting combinations and lead to an increase in perceived saturation that makes fruit objects more appealing and convincing.

Ice hockey spectators use contextual cues to guide predictive eye movements.

Eye movements are an integral part of human visual perception. They allow us to have a small foveal region with exquisite acuity and at the same time a large visual field. For a long time, eye movements were regarded as machine-like behaviors in response to visual stimulation1, but over the past few decades it has been convincingly shown that expectations, intended actions, rewards and many other cognitive factors can have profound effects on the way we move our eyes2-4. In order to be useful, our oculomotor system must minimize delay with respect to the dynamic events in the visual scene. The ability to do so has been demonstrated in situations where we are in control of these events, for example when we are making a sandwich or tea5, and when we are active participants, for example when hitting a cricket ball6. But what about scenes with complex dynamics that we do not control or directly take part in, like a hockey game we are watching as a spectator? A semantic influence on gaze fixation location during viewing of tennis videos has been suggested before7. Here we use carefully annotated hockey videos to show that the brain is indeed able to exploit the semantic context of the game to anticipate the continuous motion of the puck, leading to eye movements that are fundamentally different than when following exactly the same motion without any context.

Age effects on saccadic suppression of luminance and color.

Saccadic eye movements modulate visual perception: they initiate and terminate high acuity vision at a certain location in space, but before and during their execution visual contrast sensitivity is strongly attenuated for 100 to 200 ms. Transient perisaccadic perceptual distortions are assumed to be an important mechanism to maintain visual stability. Little is known about age effects on saccadic suppression, even though for healthy adults other major age-related changes are well documented, like a decrease of visual contrast sensitivity for intermediate and high spatial frequencies or an increase of saccade latencies. Here, we tested saccadic suppression of luminance and isoluminant chromatic flashes in 100 participants from eight to 78 years. To estimate the effect of saccadic suppression on contrast sensitivity, we used a two-alternative forced choice (2AFC) design and an adaptive staircase procedure to modulate the luminance or chromatic contrast of a flashed detection target during fixation and 15 ms after saccade onset. The target was a single horizontal luminance or chromatic line flashed 2° above or below the fixation or saccade target. Compared to fixation, average perisaccadic contrast sensitivity decreased significantly by 66% for luminance and by 36% for color. A significant correlation was found for the strength of saccadic suppression of luminance and color. However, a small age effect was found only for the strength of saccadic suppression of luminance, which increased from 64% to 70% from young to old age. We conclude that saccadic suppression for luminance and color is present in most participants independent of their age and that mechanisms of suppression stay relatively stable during healthy aging.

From Gaussian blobs to naturalistic videos: Comparison of oculomotor behavior across different stimulus complexities.

Research on eye movements has primarily been performed in two distinct ways: (1) under highly controlled conditions using simple stimuli such as dots on a uniform background, or (2) under free-viewing conditions with complex images, real-world movies, or even with observers moving around in the world. Although both approaches offer important insights, the generalizability among eye movement behaviors observed under these different conditions is unclear. Here, we compared eye movement responses to video clips showing moving objects within their natural context with responses to simple Gaussian blobs on a blank screen. Importantly, for both conditions, the targets moved along the same trajectories at the same speed. We measured standard oculometric measures for both stimulus complexities, as well as the effect of the relative angle between saccades and pursuit, and compared them across conditions. In general, eye movement responses were qualitatively similar, especially with respect to pursuit gain. For both types of stimuli, the accuracy of saccades and subsequent pursuit was highest when both eye movements were collinear. We also found interesting differences; for example, latencies of initial saccades to moving Gaussian blob targets were significantly faster compared to saccades to moving objects in video scenes, whereas pursuit accuracy was significantly higher in video scenes. These findings suggest a lower processing demand for simple target conditions during saccade preparation and an advantage for tracking behavior in natural scenes due to higher predictability provided by the context information.

Kandinsky or Me? How Free Is the Eye of the Beholder in Abstract Art?

We investigated in "art-naïve" German and Chinese participants the perception of color and spatial balance in abstract art. For color perception, we asked participants (a) to adjust the color of a single element in 24 paintings according to their liking and (b) to indicate whether they preferred their version of the painting or the original. For spatial perception, we asked participants (a) to determine the "balance point" of an artwork and (b) to indicate their preferences for the original or left-right reversed orientation of previously seen and unfamiliar paintings. Results of the color experiments suggest that, even though the interactive task was of a rather open-ended nature, observers' color adjustments were not random but systematically influenced by each painting's color palette. Overall, participants liked their own color choices about as much as the original composition. Results of the spatial experiments reveal a remarkable consistency between participants in their balance point settings. The perceived lateral position of the balance point systematically affected the left-right orientation preference for a given painting. We conclude that "art-naïve" observers are sensitive to the composition of colors and spatial structures in abstract art and are influenced by their cultural backgrounds when experiencing abstract paintings.

Dynamic combination of position and motion information when tracking moving targets.

To accurately foveate a moving target, the oculomotor system needs to estimate the position of the target at the saccade end, based on information about its position and ongoing movement, while accounting for neuronal delays and execution time of the saccade. We investigated human interceptive saccades and pursuit responses to moving targets defined by high and low luminance contrast or by chromatic contrast only (isoluminance). We used step-ramps with perpendicular directions between vertical target steps of 10 deg/s and horizontal ramps of 2.5 to 20 deg/s to separate errors with respect to the position step of the target in the vertical dimension, and errors related to target motion in the horizontal dimension. Interceptive saccades to targets of high and low luminance contrast landed close to the actual target positions, suggesting relatively accurate estimates of the amount of target displacement. Interceptive saccades to isoluminant targets were less accurate. They landed at positions the target had on average 100 ms before saccade onset. One account of this finding is that the integration of target motion is compromised for isoluminant targets moving in the periphery. In this case, the oculomotor system can use an accurate, but delayed position component, but cannot account for target movement. This deficit was also present for the postsaccadic pursuit speed. For the two luminance conditions, pursuit direction and speed were adjusted depending on the saccadic landing position. The rapid postsaccadic pursuit adjustments suggest shared position- and motion-related signals of target and eye for saccade and pursuit control.

Execution of saccadic eye movements affects speed perception.

Due to the foveal organization of our visual system we have to constantly move our eyes to gain precise information about our environment. Doing so massively alters the retinal input. This is problematic for the perception of moving objects, because physical motion and retinal motion become decoupled and the brain has to discount the eye movements to recover the speed of moving objects. Two different types of eye movements, pursuit and saccades, are combined for tracking. We investigated how the way we track moving targets can affect the perceived target speed. We found that the execution of corrective saccades during pursuit initiation modifies how fast the target is perceived compared with pure pursuit. When participants executed a forward (catch-up) saccade they perceived the target to be moving faster. When they executed a backward saccade they perceived the target to be moving more slowly. Variations in pursuit velocity without corrective saccades did not affect perceptual judgments. We present a model for these effects, assuming that the eye velocity signal for small corrective saccades gets integrated with the retinal velocity signal during pursuit. In our model, the execution of corrective saccades modulates the integration of these two signals by giving less weight to the retinal information around the time of corrective saccades.

Comparison of the precision of smooth pursuit in humans and head unrestrained monkeys.

Direct comparison of results of humans and monkeys is often complicated by differences in experimental conditions. We replicated in head unrestrained macaques experiments of a recent study comparing human directional precision during smooth pursuit eye movements (SPEM) and saccades to moving targets (Braun & Gegenfurtner, 2016). Directional precision of human SPEM follows an exponential decay function reaching optimal values of 1.5°-3° within 300 ms after target motion onset, whereas precision of initial saccades to moving targets is slightly better. As in humans, we found general agreement in the devel-opment of directional precision of SPEM over time and in the differences between direc-tional precision of initial saccades and SPEM initiation. However, monkeys showed over-all lower precision in SPEM compared to humans. This was most likely due to differences in experimental conditions, such as in the stabilization of the head, which was by a chin and a head rest in human subjects and unrestrained in monkeys.

Visual sensitivity for luminance and chromatic stimuli during the execution of smooth pursuit and saccadic eye movements.

Visual sensitivity is dynamically modulated by eye movements. During saccadic eye movements, sensitivity is reduced selectively for low-spatial frequency luminance stimuli and largely unaffected for high-spatial frequency luminance and chromatic stimuli (Nature 371 (1994), 511-513). During smooth pursuit eye movements, sensitivity for low-spatial frequency luminance stimuli is moderately reduced while sensitivity for chromatic and high-spatial frequency luminance stimuli is even increased (Nature Neuroscience, 11 (2008), 1211-1216). Since these effects are at least partly of different polarity, we investigated the combined effects of saccades and smooth pursuit on visual sensitivity. For the time course of chromatic sensitivity, we found that detection rates increased slightly around pursuit onset. During saccades to static and moving targets, detection rates dropped briefly before the saccade and reached a minimum at saccade onset. This reduction of chromatic sensitivity was present whenever a saccade was executed and it was not modified by subsequent pursuit. We also measured contrast sensitivity for flashed high- and low-spatial frequency luminance and chromatic stimuli during saccades and pursuit. During saccades, the reduction of contrast sensitivity was strongest for low-spatial frequency luminance stimuli (about 90%). However, a significant reduction was also present for chromatic stimuli (about 58%). Chromatic sensitivity was increased during smooth pursuit (about 12%). These results suggest that the modulation of visual sensitivity during saccades and smooth pursuit is more complex than previously assumed.

Discrimination of curvature from motion during smooth pursuit eye movements and fixation.

Smooth pursuit and motion perception have mainly been investigated with stimuli moving along linear trajectories. Here we studied the quality of pursuit movements to curved motion trajectories in human observers and examined whether the pursuit responses would be sensitive enough to discriminate various degrees of curvature. In a two-interval forced-choice task subjects pursued a Gaussian blob moving along a curved trajectory and then indicated in which interval the curve was flatter. We also measured discrimination thresholds for the same curvatures during fixation. Motion curvature had some specific effects on smooth pursuit properties: trajectories with larger amounts of curvature elicited lower open-loop acceleration, lower pursuit gain, and larger catch-up saccades compared with less curved trajectories. Initially, target motion curvatures were underestimated; however, ∼300 ms after pursuit onset pursuit responses closely matched the actual curved trajectory. We calculated perceptual thresholds for curvature discrimination, which were on the order of 1.5 degrees of visual angle (°) for a 7.9° curvature standard. Oculometric sensitivity to curvature discrimination based on the whole pursuit trajectory was quite similar to perceptual performance. Oculometric thresholds based on smaller time windows were higher. Thus smooth pursuit can quite accurately follow moving targets with curved trajectories, but temporal integration over longer periods is necessary to reach perceptual thresholds for curvature discrimination.NEW & NOTEWORTHY Even though motion trajectories in the real world are frequently curved, most studies of smooth pursuit and motion perception have investigated linear motion. We show that pursuit initially underestimates the curvature of target motion and is able to reproduce the target curvature ∼300 ms after pursuit onset. Temporal integration of target motion over longer periods is necessary for pursuit to reach the level of precision found in perceptual discrimination of curvature.

Dynamics of oculomotor direction discrimination.

Successful foveation of a dynamic target depends on good predictions of its movement direction and speed. We measured and compared the temporal dynamics of directional precision of both saccades and smooth pursuit and their interactions. We also compared the directional precision of both eye movements to psychophysical direction discrimination thresholds. Directional thresholds of pure pursuit responses improved rapidly and reached asymptotic values of 1.5°-3° within 300 ms after target motion onset, both for trained and untrained observers and irrespective of the speed of the stimuli. Psychophysical thresholds were in the same range. Directional thresholds for saccades in the ramp paradigm were just slightly higher, but these occurred significantly earlier in time at around 200 ms after target motion onset. At the equivalent time during pure pursuit initiation, thresholds were typically higher by 2°-3°. The rise in directional precision-or decrease in thresholds-over time was more pronounced for trials with longer latencies. As an effect, precision depended mainly on time since stimulus motion onset rather than pursuit onset. Directional precision for saccades to static targets was slightly better than to moving targets, at even shorter latencies. We conclude that directional precision is higher for the saccadic system at saccade onset than for the pursuit system, presumably due to additional position signals that are not available to the pursuit system at that point in time. The pursuit response improves rapidly due to refined sensory processing and motor planning. The combination of initial saccades and pursuit to track moving targets is a good strategy for the oculomotor system to reduce directional errors during the phase of initiation. The target speed has very little effects on the directional precision of both eye movements.

Eye movements and perception: a selective review.

Eye movements are an integral and essential part of our human foveated vision system. Here, we review recent work on voluntary eye movements, with an emphasis on the last decade. More selectively, we address two of the most important questions about saccadic and smooth pursuit eye movements in natural vision. First, why do we saccade to where we do? We argue that, like for many other aspects of vision, several different circuits related to salience, object recognition, actions, and value ultimately interact to determine gaze behavior. Second, how are pursuit eye movements and perceptual experience of visual motion related? We show that motion perception and pursuit have a lot in common, but they also have separate noise sources that can lead to dissociations between them. We emphasize the point that pursuit actively modulates visual perception and that it can provide valuable information for motion perception.

Challenges to normal neural functioning provide insights into separability of motion processing mechanisms.

There is a long history of attempts to disentangle different visual processing mechanisms for physically different motion cues. However, underlying neural correlates and separability of networks are still under debate. We aimed to refine the current understanding by studying differential vulnerabilities when normal neural functioning is challenged. We investigated effects of ageing and extrastriate brain lesions on detection thresholds for motion defined by either luminance- or contrast modulations, known as first- and second-order motion. Both approaches focus on extrastriate processing changes and combine distributed as well as more focal constraints. Our ageing sample comprised 102 subjects covering an age range from 20 to 82 years. Threshold signal-to-noise ratios for detection approximately doubled across the age range for both motion types. Results suggest that ageing affects perception of both motion types to an equivalent degree and thus support overlapping processing resources. Underlying neural substrates were further qualified by testing perceptual performance of 18 patients with focal cortical brain lesions. We determined selective first-order motion deficits in three patients, selective second-order motion deficits in only one patient, and deficits for both motion types in three patients. Lesion analysis yielded support for common functional substrates in higher cortical regions. Functionally specific substrates remained ambiguous, but tended to cover earlier visual areas. We conclude that observed vulnerabilities of first- and second-order motion perception provide limited evidence for functional specialization at early extrastriate stages, but emphasize shared processing pathways at higher cortical levels.

Keep your eyes on the ball: smooth pursuit eye movements enhance prediction of visual motion.

Success of motor behavior often depends on the ability to predict the path of moving objects. Here we asked whether tracking a visual object with smooth pursuit eye movements helps to predict its motion direction. We developed a paradigm, "eye soccer," in which observers had to either track or fixate a visual target (ball) and judge whether it would have hit or missed a stationary vertical line segment (goal). Ball and goal were presented briefly for 100-500 ms and disappeared from the screen together before the perceptual judgment was prompted. In pursuit conditions, the ball moved towards the goal; in fixation conditions, the goal moved towards the stationary ball, resulting in similar retinal stimulation during pursuit and fixation. We also tested the condition in which the goal was fixated and the ball moved. Motion direction prediction was significantly better in pursuit than in fixation trials, regardless of whether ball or goal served as fixation target. In both fixation and pursuit trials, prediction performance was better when eye movements were accurate. Performance also increased with shorter ball-goal distance and longer presentation duration. A longer trajectory did not affect performance. During pursuit, an efference copy signal might provide additional motion information, leading to the advantage in motion prediction.

Does the noise matter? Effects of different kinematogram types on smooth pursuit eye movements and perception.

We investigated how the human visual system and the pursuit system react to visual motion noise. We presented three different types of random-dot kinematograms at five different coherence levels. For transparent motion, the signal and noise labels on each dot were preserved throughout each trial, and noise dots moved with the same speed as the signal dots but in fixed random directions. For white noise motion, every 20 ms the signal and noise labels were randomly assigned to each dot and noise dots appeared at random positions. For Brownian motion, signal and noise labels were also randomly assigned, but the noise dots moved at the signal speed in a direction that varied randomly from moment to moment. Neither pursuit latency nor early eye acceleration differed among the different types of kinematograms. Late acceleration, pursuit gain, and perceived speed all depended on kinematogram type, with good agreement between pursuit gain and perceived speed. For transparent motion, pursuit gain and perceived speed were independent of coherence level. For white and Brownian motions, pursuit gain and perceived speed increased with coherence but were higher for white than for Brownian motion. This suggests that under our conditions, the pursuit system integrates across all directions of motion but not across all speeds.

Localization of speed differences of context stimuli during fixation and smooth pursuit eye movements.

The visual system can detect speed changes of moving objects only by means of alterations of retinal image motion, which is also subject to changes induced by head or eye movements. Here we investigated whether smooth pursuit eye movements affect the ability to localize short speed perturbations of large context stimuli. Psychophysical thresholds for localization, discrimination and detection of speed perturbations in one of two context stimuli were measured under two main conditions: in fixation trials subjects fixated a central stationary spot, in pursuit trials they followed a horizontally moving target with their eyes. Context stimuli were vertically oriented sine wave gratings moving simultaneously above and below the fixation or pursuit target for one second in the same direction at the same or a different speed as the pursuit target. During the movement one of the gratings suddenly changed its speed for 500 ms and returned to its original speed. Observers were asked to discern the location of the speed change (two-alternative spatial forced choice task). While detection (two-interval forced choice) and discrimination thresholds for the kind of speed perturbation were in the normal range of Weber fractions of 10-15%, thresholds for the location of the speed perturbation were dramatically increased to 30-50%. Localization thresholds were particularly high when the retinal motion was mainly due to the context movements as during fixation or slow pursuit and significantly reduced when the retinal motion was mainly due to pursuit. This result indicates that the origin of retinal motion, whether it is caused by object motion or by voluntary pursuit is important. We conclude that the localization of speed perturbations affecting one of two peripheral moving objects is exceedingly complicated for the visual system probably due to the dominance of relative motion. During smooth pursuit the ability to localize speed perturbations of non-foveated objects seems to be improved by additional information gained from pursuit such as corollary discharge.

Improved visual sensitivity during smooth pursuit eye movements: temporal and spatial characteristics.

Recently, we showed that contrast sensitivity for color and high-spatial frequency luminance stimuli is enhanced during smooth pursuit eye movements (Schütz et al., 2008). In this study, we investigated the enhancement over a wide range of temporal and spatial frequencies. In Experiment 1, we measured the temporal impulse response function (TIRF) for colored stimuli. The TIRF for pursuit and fixation differed mostly with respect to the gain but not with respect to the natural temporal frequency. Hence, the sensitivity enhancement seems to be rather independent of the temporal frequency of the stimuli. In Experiment 2, we measured the spatial contrast sensitivity function for luminance-defined Gabor patches with spatial frequencies ranging from 0.2 to 7 cpd. We found a sensitivity improvement during pursuit for spatial frequencies above 2-3 cpd. Between 0.5 and 3 cpd, sensitivity was impaired by smooth pursuit eye movements, but no consistent difference was observed below 0.5 cpd. The results of both experiments are consistent with an increased contrast gain of the parvocellular retinogeniculate pathway.

Object recognition during foveating eye movements.

We studied how saccadic and smooth pursuit eye movements affect the recognition of briefly presented letters appearing within the eye movement target. First we compared the recognition performance during steady-state pursuit and during fixation. Single letters were presented for seven different durations ranging from 10 to 400 ms and four contrast levels ranging from 5% to 40%. For both types of eye movements the recognition rates increased with duration and contrast, but they were on average 11% lower during pursuit. In daily life humans use a combination of saccadic and smooth pursuit eye movements to foveate a peripheral moving object. To investigate this more natural situation, we presented a peripheral target that was either stationary or moving horizontally, above or below the fixation spot. Participants were asked to saccade to the target and to keep it foveated. The letters were presented at different times relative to the first target directed saccade. As would be expected from retinal masking and motion blur during saccades, the discrimination performance increased with increasing post-saccadic delay. If the target moved and the saccade was followed by pursuit, letter recognition performance was on average 16% lower than if the target was stationary and the saccade was followed by fixation.

Cortical networks for motion processing: effects of focal brain lesions on perception of different motion types.

Neuropsychological studies in humans provide evidence for a variety of extrastriate cortical areas involved in visual motion perception. Multiple mechanisms underlying processing of different motion types have been proposed, however, support for cortical specialization has remained controversial so far. We therefore studied motion perception in 23 patients with focal lesions to various cortical areas and considered translational motion, heading from radial flow, as well as biological motion. Patients' detection thresholds were compared with age-specific data from a large healthy control sample (n=122). Elevated thresholds and significant threshold asymmetries between both visual hemifields were defined as deficits. Contrary to prevalent opinion, we found a high prevalence of motion deficits in our sample. Impairment was restricted to a specific motion type in 10 patients, whereas only a single patient showed a deficit for multiple motion types. Functional areas were determined by lesion density plots and by comparison between patients with and without a specific deficit. Results emphasize a dissociation between basic motion processing and processing of complex motion. Anatomical analysis confirmed critical occipito-temporo-parietal areas for perception of translational motion. In contrast, heading perception from radial flow proved to be remarkably robust to most lesions. We exclusively identified the frontal eye fields as a critical structure. Biological motion perception relied on distinct pathways involving temporal, parietal, and frontal areas. Although precise functional roles of identified areas cannot be determined conclusively, results clearly indicate regional specialization for motion types of different complexity. We propose a network for motion processing involving widely distributed cortical areas.