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In natural environments, head movements are required to search for objects outside the field of view (FoV). Here we investigate the power of a salient target in an extended visual search array to facilitate faster detection once this item comes into the FoV by a head movement. We conducted two virtual reality experiments using spatially clustered sets of stimuli to observe target detection and head and eye movements during visual search. Participants completed search tasks with three conditions: (1) target in the initial FoV, (2) head movement needed to bring the target into the FoV, (3) same as condition 2 but the periphery was initially hidden and appeared after the head movement had brought the location of the target set into the FoV. We measured search time until participants found a more salient (O) or less salient (T) target among distractors (L). On average O's were found faster than T's. Gaze analysis showed that saliency facilitation occurred due to the target guiding the search only if it was within the initial FoV. When targets required a head movement to enter the FoV, participants followed the same search strategy as in trials without a visible target in the periphery. Moreover, faster search times for salient targets were only caused by the time required to find the target once the target set was reached. This suggests that the effect of stimulus saliency differs between visual search on fixed displays and when we are actively searching through an extended visual field., (© 2024. The Author(s).)

Self-motion generates optic flow, a pattern of expanding visual motion. Heading estimation from optic flow analysis is accurate in rigid environments, but it becomes challenging when other human walkers introduce independent motion to the scene. Previous studies showed that heading perception is surprisingly accurate when moving through a crowd of walkers but revealed strong heading biases when either articulation or translation of biological motion were presented in isolation. We hypothesized that these biases resulted from misperceiving the self-motion as curvilinear. Such errors might manifest as opposite biases depending on whether the observer perceived the crowd motion as indication of his/her self-translation or self-rotation. Our study investigated the link between heading biases and illusory path perception. Participants assessed heading and path perception while observing optic flow stimuli with varying walker movements. Self-motion perception was accurate during natural locomotion (articulation and translation), but significant heading biases occurred when walkers only articulated or translated. In this case, participants often reported a curved path of travel. Heading error and curvature pointed in opposite directions. On average, participants perceived the walker motion as evidence for viewpoint rotation leading to curvilinear path percepts.

Vision requires that we rotate our eyes frequently to look at informative structures in the scene. Eye movements are planned by the brain but their execution depends on the mechanical properties of the oculomotor plant, that is, the arrangement of eyeball position, muscle insertions, and pulley locations. Therefore, the biomechanics of rotations is sensitive to eyeball translation because it changes muscle levers. Eyeball translations are little researched as they are difficult to measure with conventional techniques. Here, we investigated the effects of eyeball translation on the coordination of eyeball rotation by high-speed MRI recordings of saccadic eye movements during blinks, which are known to produce strong translations. We found that saccades during blinks massively overshoot their targets and that these overshoots occur in a transient fashion such that the gaze is back on target at the time the blink ends. These dynamic overshoots were tightly coupled to the eyeball translation, both in time and in size. Saccades made without blinks were also accompanied by small amounts of transient eyeball retraction, the size of which scaled with saccade amplitude. These findings demonstrate a complex combination of rotation and translation of the eye. The mechanical consequences of eyeball translation on oculomotor control should be considered along with the neural implementation in the brainstem to understand the generation of eye movements and their disorders. NEW & NOTEWORTHY We found that saccades during blinks can massively overshoot their target when the eyeball is retracted. Our data imply that the overshoots are not part of the saccade plan prepared in the brainstem, but instead a consequence of the altered biomechanics resulting from concurrent eyeball translation and rotation. To our best knowledge, this is the first direct observation of dynamic properties of the oculomotor plant altering the execution of rotational eye movements.

The blind spot is both a necessity and a nuisance for seeing. It is the portion of the visual field projecting to where the optic nerve crosses the retina, a region devoid of photoreceptors and hence visual input. The precise way in which vision transitions into blindness at the blind spot border is to date unknown. A chief challenge to map this transition is the incessant movement of the eye, which unavoidably smears measurements across space. In this study, we used high-resolution eye-tracking and state-of-the-art retinal stabilization to finely map the blind spot borders. Participants reported the onset of tiny high-contrast probes that were briefly flashed at precise positions around the blind spot. This method has sufficient resolution to enable mapping of blood vessels from psychophysical measurements. Our data show that, even after accounting for eye movements, the transition zones at the edges of the blind spot are considerable. On the horizontal meridian, the regions with detection rates between 80% and 20% span approximately 25% of the overall width of the blind spot. These borders also vary considerably in size across different axes. These data show that the transition from full visibility to blindness at the blind spot border is not abrupt but occurs over a broad area., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023. Published by Elsevier Ltd.)

Humans can use visual motion to estimate the distance they have traveled. In static environments, optic flow generated by self-motion provides a pattern of expanding motion that is used for the estimation of travel distance. When the environment is populated by other people, their biological motion destroys the one-to-on correspondence between optic flow and travel distance. We investigated how observers estimate travel distance in a crowded environment. In three conditions, we simulated self-motion through a crowd of standing, approaching, or leading point-light walkers. For a standing crowd, optic flow is a veridical signal for distance perception. For an approaching crowd, the visual motion is the sum of the self-motion-induced optic flow and the optic flow produced by the approaching walkers. If only optic flow were to be used, travel distance estimates would be too high because of the approaching direction of the crowd toward the observer. If, on the other hand, cues from biological motion could be used to estimate the speed of the crowd, then the excessive optic from the approaching crowd flow might be compensated. In the leading crowd condition, in which walkers of the crowd keep their distance from the observer as they walk along with the observer, no optic flow is produced. In this condition, travel distance estimation would have to rely solely on biological motion information. We found that distance estimation was quite similar across these three conditions. This suggests that biological motion information can be used (a) to compensate for excessive optic flow in the approaching crowd condition and (b) to generate distance information in the leading crowd condition.

Visual selection is characterized by a trade-off between speed and accuracy. Speed or accuracy of the selection process can be affected by higher level factors-for example, expecting a reward, obtaining task-relevant information, or seeing an intrinsically relevant target. Recently, motivation by reward has been shown to simultaneously increase speed and accuracy, thus going beyond the speed-accuracy-trade-off. Here, we compared the motivating abilities of monetary reward, task-relevance, and image content to simultaneously increase speed and accuracy. We used a saccadic distraction task that required suppressing a distractor and selecting a target. Across different blocks successful target selection was followed either by (i) a monetary reward, (ii) obtaining task-relevant information, or (iii) seeing the face of a famous person. Each block additionally contained the same number of irrelevant trials lacking these consequences, and participants were informed about the upcoming trial type. We found that postsaccadic vision of a face affected neither speed nor accuracy, suggesting that image content does not affect visual selection via motivational mechanisms. Task relevance increased speed but decreased selection accuracy, an observation compatible with a classical speed-accuracy trade-off. Motivation by reward, however, simultaneously increased response speed and accuracy. Saccades in all conditions deviated away from the distractor, suggesting that the distractor was suppressed, and this deviation was strongest in the reward block. Drift-diffusion modelling revealed that task-relevance affected behavior by affecting decision thresholds, whereas motivation by reward additionally increased the rate of information uptake. The present findings thus show that the three consequences differ in their motivational abilities., (© 2022. The Author(s).)

Binding theories assume that stimulus and response features are integrated into short-lasting episodes and that upon repetition of any feature the whole episode is retrieved, thereby affecting performance. Such binding theories are nowadays the standard explanation for a wide range of action control tasks and aim to explain all simple actions, without making assumptions of effector specificity. Yet, it is unclear if eye movements are affected by integration and retrieval in the same way as manual responses. We asked participants to discriminate letters framed by irrelevant shapes. In Experiment 1, participants gave their responses with eye movements. Saccade landing positions showed a spatial error pattern consistent with predictions of binding theories. Saccadic latencies were not affected. In Experiment 2 with an increased interval between prime and probe, the error pattern diminished, again congruent with predictions of binding theories presuming quickly decaying retrieval effects. Experiment 3 used the same task as in Experiment 1, but participants executed their responses with manual key presses; again, we found a binding pattern in response accuracy. We conclude that eye movements and manual responses are affected by the same integration and retrieval processes, supporting the tacit assumption of binding theories to apply to any effector., (© 2022. The Author(s).)

Saccadic adaptation ($SA$) is a cerebellar-dependent learning of motor commands ($MC$), which aims at preserving saccade accuracy. Since $SA$ alters visual localization during fixation and even more so across saccades, it could also involve changes of target and/or saccade visuospatial representations, the latter ($CDv$) resulting from a motor-to-visual transformation (forward dynamics model) of the corollary discharge of the $MC$. In the present study, we investigated if, in addition to its established role in adaptive adjustment of $MC$, the cerebellum could contribute to the adaptation-associated perceptual changes. Transfer of backward and forward adaptation to spatial perceptual performance (during ocular fixation and trans-saccadically) was assessed in eight cerebellar patients and eight healthy volunteers. In healthy participants, both types of $SA$ altered $MC$ as well as internal representations of the saccade target and of the saccadic eye displacement. In patients, adaptation-related adjustments of $MC$ and adaptation transfer to localization were strongly reduced relative to healthy participants, unraveling abnormal adaptation-related changes of target and $CDv$. Importantly, the estimated changes of $CDv$ were totally abolished following forward session but mainly preserved in backward session, suggesting that an internal model ensuring trans-saccadic localization could be located in the adaptation-related cerebellar networks or in downstream networks, respectively., (© The Author(s) 2021. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Humans visually inspect the world with their fovea and select new parts of the scene using saccadic eye movements. Foveal inspection and the decision of where and when to look next proceed simultaneously, but there is mixed evidence concerning their independence. Here, we tested their interdependence using drift-diffusion modeling. Participants first made a saccade to a predetermined inspection target and subsequently decided between two selection targets. We found that the inspected target's meaningfulness and the opportunity to preview it peripherally affects fixation durations and the upcoming saccadic selection. Drift-diffusion modeling showed that meaningfulness and the absence of peripheral preview can both delay the subsequent saccadic decision process and affect the rate at which peripheral information is accumulated. Our results thus show that foveal inspection and peripheral selection are dependent on each other and that peripheral information can be maintained across the saccade to influence subsequent eye movement decisions., Competing Interests: The authors declare no competing interests., (© 2022 The Author(s).)

Saccadic eye movements are often imprecise and result in an error between expected and actual retinal target location after the saccade. Repeated experience of this error produces changes in saccade amplitude to reduce the error and concomitant changes in apparent visual location. We investigated the relationship between these two plastic processes in a series of experiments. Following a recent paradigm of inhibition of saccadic adaptation, in which participants are instructed to look at the initial target position and to continue to look at that position even if the target were to move again, our participants nevertheless perceived a visual probe presented near the saccade target to be shifted in direction of the target error. The location percept of the target gradually shifted and diverged over time from the executed saccade. Our findings indicate that changes in perceived location can be the same even when changes in saccade amplitude differ according to instruction and can develop even when the amplitude of the saccades executed during the adaptation procedure does not change. There are two possible explanations for this divergence between the adaptation states of saccade amplitude and perceived location. Either the intrasaccadic target step might trigger updating of the association between pre- and post-saccadic target positions, causing the localization shift, or the saccade motor command adjusts together with the perceived location at a common adaptation site, downstream from which voluntary control is exerted upon the executed eye movement only.

Nonrigid materials such as liquids or smoke deform over time. Little is known about the visual perception of nonrigid motion other than that many motion cues associated with rigid motion perception are not reliable for nonrigid motion. Nonrigid motion patterns lack clear borders and their movement can be inconsistent with the motion of their parts. We developed a novel stimulus that creates a nonrigid vortex motion pattern in a random dot distribution and decouples the movement of the vortex from the first-order motion of the dots. We presented three moving vortices that entailed consecutively fewer motion cues, eliminating occlusion, motion borders, and velocity field gradients in the process. Subjects were well able to report the end position and travel path in all cases, showing that nonrigid motion is perceived through an analysis of the temporal evolution of visual motion patterns and does not require borders or speed differences. Adding a coherent global motion did not hamper perception, but adding local noise did, indicating that the visual system uses mid-level features that are on a local scale. We also found that participants judged the movement of the nonrigid motion patterns slower than a rigid control, revealing that speed perception was based on a combination of motion of the parts and movement of the pattern. We propose that the visual system uses the temporal evolution of a motion pattern for the perception of nonrigid motion and suggest a plausible mechanism based on the curl of the motion field., (© The Author(s) 2022. Published by Oxford University Press on behalf of the National Academy of Sciences.)

Our eyes are constantly in motion and the various kinds of eye movements are closely linked to many aspects of human cognitive processing. Measuring all possible eye movements unobtrusively is not achievable with current methods. Video-based eye-trackers only measure rotational but not translational motion of the eye, require a calibration process relying on the participant's self-report of accurate fixation, and do not work if vision of the eyeball is blocked. Scleral search coils attach physical weight on the eyeball and also do not measure translation. Here, we describe a novel and fully automated method to use real-time magnetic resonance imaging (MRI) for eye tracking. We achieved a temporal resolution sufficient to measure eye rotations and translations as short as those that occur within a blink and behind a closed eyelid. To demonstrate this method, we measured the full extent of the blink-related eye movement for two individuals, suggesting that the eye approaches a holding position during lid closure and can move by as much as 35° in rotation and 2 mm in translation. We also investigated the coordination of gaze shifts with blinks. We found that the gaze shift is tightly coupled in time to the translational blink movement and that blinks can induce significant temporal shifts of the gaze trajectory between left and right eye. Our MR-based Eye Tracking (MREyeTrack) method allows measurement of eye movements in terms of both translation and rotation and enables new opportunities for studying ocular motility and its disorders., (Copyright © 2022 Kirchner et al.)

Saccadic eye movements bring objects of interest onto our fovea. These gaze shifts are essential for visual perception of our environment and the interaction with the objects within it. They precede our actions and are thus modulated by current goals. It is assumed that saccadic adaptation, a recalibration process that restores saccade accuracy in case of error, is mainly based on an implicit comparison of expected and actual post-saccadic position of the target on the retina. However, there is increasing evidence that task demands modulate saccade adaptation and that errors in task performance may be sufficient to induce changes to saccade amplitude. We investigated if human participants are able to flexibly use different information sources within the post-saccadic visual feedback in task-dependent fashion. Using intra-saccadic manipulation of the visual input, participants were either presented with congruent post-saccadic information, indicating the saccade target unambiguously, or incongruent post-saccadic information, creating conflict between two possible target objects. Using different task instructions, we found that participants were able to modify their saccade behavior such that they achieved the goal of the task. They succeeded in decreasing saccade gain or maintaining it, depending on what was necessary for the task, irrespective of whether the post-saccadic feedback was congruent or incongruent. It appears that action intentions prime task-relevant feature dimensions and thereby facilitated the selection of the relevant information within the post-saccadic image. Thus, participants use post-saccadic feedback flexibly, depending on their intentions and pending actions.

Progressive addition lenses (PALs) are ophthalmic lenses to correct presbyopia by providing improvements of near and far vision in different areas of the lens, but distorting the periphery of the wearer's field of view. Distortion-related difficulties reported by PAL wearers include unnatural self-motion perception. Visual self-motion perception is guided by optic flow, the pattern of retinal motion produced by self-motion. We tested the influence of PAL distortions on optic flow-based heading estimation using a model of heading perception and a virtual reality-based psychophysical experiment. The model predicted changes of heading estimation along a vertical axis, depending on visual field size and gaze direction. Consistent with this prediction, participants experienced upwards deviations of self-motion when gaze through the periphery of the lens was simulated, but not for gaze through the center. We conclude that PALs may lead to illusions of self-motion which could be remedied by a careful gaze strategy., Competing Interests: The authors declare that there are no conflicts of interest that could have interfered in the course of this study., (© 2021 The Author(s).)

Short-latency saccades are often biased toward salient objects or toward the center of images, for example, when inspecting photographs of natural scenes. Here, we measured the contribution of salient objects and central fixation bias to visual selection over time. Participants made saccades to images containing one salient object on a structured background and were instructed to either look at (i) the image center, (ii) the salient object, or (iii) at a cued position halfway in between the two. Results revealed, first, an early involuntary bias toward the image center irrespective of strategic behavior or the location of objects in the image. Second, the salient object bias was stronger than the center bias and prevailed over the latter when they directly competed for visual selection. In a second experiment, we tested whether the center bias depends on how well the image can be segregated from the monitor background. We asked participants to explore images that either did or did not contain a salient object while we manipulated the contrast between image background and monitor background to make the image borders more or less visible. The initial orienting toward the image was not affected by the image-monitor contrast, but only by the presence of objects-with a strong bias toward the center of images containing no object. Yet, a low image-monitor contrast reduced this center bias during the subsequent image exploration.

Humans and other primates are equipped with a foveated visual system. As a consequence, we reorient our fovea to objects and targets in the visual field that are conspicuous or that we consider relevant or worth looking at. These reorientations are achieved by means of saccadic eye movements. Where we saccade to depends on various low-level factors such as a targets' luminance but also crucially on high-level factors like the expected reward or a targets' relevance for perception and subsequent behavior. Here, we review recent findings how the control of saccadic eye movements is influenced by higher-level cognitive processes. We first describe the pathways by which cognitive contributions can influence the neural oculomotor circuit. Second, we summarize what saccade parameters reveal about cognitive mechanisms, particularly saccade latencies, saccade kinematics and changes in saccade gain. Finally, we review findings on what renders a saccade target valuable, as reflected in oculomotor behavior. We emphasize that foveal vision of the target after the saccade can constitute an internal reward for the visual system and that this is reflected in oculomotor dynamics that serve to quickly and accurately provide detailed foveal vision of relevant targets in the visual field., (© The Author(s) 2021.)

During locomotion, leg movements define the direction of walking (forward or backward) and the path one is taking (straight or curved). These aspects of locomotion produce characteristic visual motion patterns during movement. Here, we tested whether cortical regions responding to either egomotion-compatible visual motion, or leg movements, or both, are sensitive to these locomotion-relevant aspects of visual motion. We compared a curved path (typically the visual feedback of a changing direction of movement in the environment) to a linear path for simulated forward and backward motion in an event-related fMRI experiment. We used an individual surface-based approach and two functional localizers to define (1) six egomotion-related areas (V6+, V3A, intraparietal motion area [IPSmot], cingulate sulcus visual area [CSv], posterior cingulate area [pCi], posterior insular cortex [PIC]) using the flow field stimulus and (2) three leg-related cortical regions (human PEc [hPEc], human PE [hPE] and primary somatosensory cortex [S-I]) using a somatomotor task. Then, we extracted the response from all these regions with respect to the main event-related fMRI experiment, consisting of passive viewing of an optic flow stimulus, simulating a forward or backward direction of self-motion in either linear or curved path. Results showed that some regions have a significant preference for the curved path motion (hPEc, hPE, S-I, IPSmot) or a preference for the forward motion (V3A), while other regions have both a significant preference for the curved path motion and for the forward compared to backward motion (V6+, CSv, pCi). We did not find any significant effects of the present stimuli in PIC. Since controlling locomotion mainly means controlling changes of walking direction in the environment during forward self-motion, such a differential functional profile among these cortical regions suggests that they play a differentiated role in the visual guidance of locomotion., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Sensorimotor learning adapts motor output to maintain movement accuracy. For saccadic eye movements, learning also alters space perception, suggesting a dissociation between the performed saccade and its internal representation derived from corollary discharge (CD). This is critical since learning is commonly believed to be driven by CD-based visual prediction error. We estimate the internal saccade representation through pre- and trans-saccadic target localization, showing that it decouples from the actual saccade during learning. We present a model that explains motor and perceptual changes by collective plasticity of spatial target percept, motor command, and a forward dynamics model that transforms CD from motor into visuospatial coordinates. We show that learning does not follow visual prediction error but instead a postdictive update of space after saccade landing. We conclude that trans-saccadic space perception guides motor learning via CD-based postdiction of motor error under the assumption of a stable world., Competing Interests: JM, ML No competing interests declared, (© 2021, Masselink and Lappe.)

Humans scan their visual environment using saccade eye movements. Where we look is influenced by bottom-up salience and top-down factors, like value. For reactive saccades in response to suddenly appearing stimuli, it has been shown that short-latency saccades are biased towards salience, and that top-down control increases with increasing latency. Here, we show, in a series of six experiments, that this transition towards top-down control is not determined by the time it takes to integrate value information into the saccade plan, but by the time it takes to inhibit suddenly appearing salient stimuli. Participants made consecutive saccades to three fixation crosses and a vertical bar consisting of a high-salient and a rewarded low-salient region. Endpoints on the bar were biased towards salience whenever it appeared or reappeared shortly before the last saccade was initiated. This was also true when the eye movement was already planned. When the location of the suddenly appearing salient region was predictable, saccades were aimed in the opposite direction to nullify this sudden onset effect. Successfully inhibiting salience, however, could only be achieved by previewing the target. These findings highlight the importance of inhibition for top-down eye-movement control.

Perceiving object motion during self-movement is an essential ability of humans. Previous studies have reported that the visual system can use both visual information (such as optic flow) and non-visual information (such as vestibular, somatosensory, and proprioceptive information) to identify and globally subtract the retinal motion component due to self-movement to recover scene-relative object motion. In this study, we used a motion-nulling method to directly measure and quantify the contribution of visual and non-visual information to the perception of scene-relative object motion during walking. We found that about 50% of the retinal motion component of the probe due to translational self-movement was removed with non-visual information alone and about 80% with visual information alone. With combined visual and non-visual information, the self-movement component was removed almost completely. Although non-visual information played an important role in the removal of self-movement-induced retinal motion, it was associated with decreased precision of probe motion estimates. We conclude that neither non-visual nor visual information alone is sufficient for the accurate perception of scene-relative object motion during walking, which instead requires the integration of both sources of information.

Heading estimation from optic flow relies on the assumption that the visual world is rigid. This assumption is violated when one moves through a crowd of people, a common and socially important situation. The motion of people in the crowd contains cues to their translation in the form of the articulation of their limbs, known as biological motion. We investigated how translation and articulation of biological motion influence heading estimation from optic flow for self-motion in a crowd. Participants had to estimate their heading during simulated self-motion toward a group of walkers who collectively walked in a single direction. We found that the natural combination of translation and articulation produces surprisingly small heading errors. In contrast, experimental conditions that either present only translation or only articulation produced strong idiosyncratic biases. The individual biases explained well the variance in the natural combination. A second experiment showed that the benefit of articulation and the bias produced by articulation were specific to biological motion. An analysis of the differences in biases between conditions and participants showed that different perceptual mechanisms contribute to heading perception in crowds. We suggest that coherent group motion affects the reference frame of heading perception from optic flow.

Heading estimation from optic flow is crucial for safe locomotion but becomes inaccurate if independent object motion is present. In ecological settings, such motion typically involves other animals or humans walking across the scene. An independently walking person presents a local disturbance of the flow field, which moves across the flow field as the walker traverses the scene. Is the bias in heading estimation produced by the local disturbance of the flow field or by the movement of the walker through the scene? We present a novel flow field stimulus in which the local flow disturbance and the movement of the walker can be pitted against each other. Each frame of this stimulus consists of a structureless random dot distribution. Across frames, the body shape of a walker is molded by presenting different flow field dynamics within and outside the body shape. In different experimental conditions, the flow within the body shape can be congruent with the walker's movement, incongruent with it, or congruent with the background flow. We show that heading inaccuracy results from the local flow disturbance rather than the movement through the scene. Moreover, we show that the local disturbances of the optic flow can be used to segment the walker and support biological motion perception to some degree. The dichotomous result that the walker can be segmented from the scene but that heading perception is nonetheless influenced by the flow produced by the walker confirms separate visual pathways for heading estimation, object segmentation, and biological motion perception.

Systematic shortening or lengthening of target objects during saccades modifies saccade amplitudes and perceived size of the objects. These two events are concomitant when size change during the saccade occurs asymmetrically, thereby shifting the center of mass of the object. In the present study, we asked whether or not the two are necessarily linked. We tested human participants in symmetrical systematic shortening and lengthening of a vertical bar during a horizontal saccade, aiming to not modify the saccade amplitude. Before and after a phase of trans-saccadic changes of the target bar, participants manually indicated the sizes of various vertically oriented bars by open-loop grip aperture. We evaluated the effect of trans-saccadic changes of bar length on manual perceptual reports and whether this change depended on saccade amplitude. As expected, we did not induce any change in horizontal or vertical components of saccade amplitude, but we found a significant difference in perceived size after the lengthening experiment compared to after the shortening experiment. Moreover, after the lengthening experiment, perceived size differed significantly from pre-lengthening baseline. These findings suggest that a change of size perception can be induced trans-saccadically, and its mechanism does not depend on saccadic amplitude change.

We investigated whether biological motion biases heading estimation from optic flow in a similar manner to nonbiological moving objects. In two experiments, observers judged their heading from displays depicting linear translation over a random-dot ground with normal point light walkers, spatially scrambled point light walkers, or laterally moving objects composed of random dots. In Experiment 1, we found that both types of walkers biased heading estimates similarly to moving objects when they obscured the focus of expansion of the background flow. In Experiment 2, we also found that walkers biased heading estimates when they did not obscure the focus of expansion. These results show that both regular and scrambled biological motion affect heading estimation in a similar manner to simple moving objects, and suggest that biological motion is not preferentially processed for the perception of self-motion.

Perisaccadic compression of the perceived location of flashed visual stimuli toward a saccade target occurs from about 50 ms before a saccade. Here we show that between 150 and 80 ms before a saccade, perceived locations are shifted toward the fixation point. To establish the cause of the "reverse" presaccadic perceptual distortion, participants completed several versions of a saccade task. After a cue to saccade, a probe bar stimulus was briefly presented within the saccade trajectory. In Experiment 1 participants made (a) overlap saccades with immediate return saccades, (b) overlap saccades, and (c) step saccades. In Experiment 2 participants made gap saccades in complete darkness. In Experiment 3 participants maintained fixation with the probe stimuli masked at various interstimulus intervals. Participants indicated the bar's location using a mouse cursor. In all conditions in Experiment 1 presaccadic compression was preceded by compression toward the initial fixation. In Experiment 2, saccadic compression was maintained but the preceding countercompression was not observed. Stimuli masked at fixation were not compressed. This suggests the two opposing compression effects are related to the act of executing an eye movement. They are also not caused by the requirement to make two sequential saccades ending at the initial fixation location and are not caused by continuous presence of the fixation markers. We propose that countercompression is related to fixation activity and is part of the sequence of motor preparations to execute a cued saccade.

We investigated whether the proprioceptive eye position signal after the execution of a saccadic eye movement is used to estimate the accuracy of the movement. If so, saccadic adaptation, the mechanism that maintains saccade accuracy, could use this signal in a similar way as it uses visual feedback after the saccade. To manipulate the availability of the proprioceptive eye position signal we utilized the finding that proprioceptive eye position information builds up gradually after a saccade over a time interval comparable to typical saccade latencies. We confined the retention time of gaze at the saccade landing point by asking participants to make fast return saccades to the fixation point that preempt the usability of proprioceptive eye position signals. In five experimental conditions we measured the influence of the visual and proprioceptive feedback, together and separately, on the development of adaptation. We found that the adaptation of the previously shortened saccades in the case of visual feedback being unavailable after the saccade was significantly weaker when the use of proprioceptive eye position information was impaired by fast return saccades. We conclude that adaptation can be driven by proprioceptive eye position feedback. NEW & NOTEWORTHY We show that proprioceptive eye position information is used after a saccade to estimate motor error and adapt saccade control. Previous studies on saccadic adaptation focused on visual feedback about saccade accuracy. A multimodal error signal combining visual and proprioceptive information is likely more robust. Moreover, combining proprioceptive and visual measures of saccade performance can be helpful to keep vision, proprioception, and motor control in alignment and produce a coherent representation of space.

Adaptation to changes in the environment allows the visual system to achieve optimal perception in a continuously changing visual world. One particular example regarding recurrently encountered changes in everyday vision is geometrical distortions of the environment when wearing spectacles for vision correction, e.g., image shear by skew geometric distortions in progressive additional lenses. For optimal visual performance, it would be beneficial if the visual system uses previous history of recurrent distortions and learns to adapt fast when they are reapplied, yet this has not been systematically shown. The present study evaluates experience-dependent long-term facilitation of fast adaptation to image skew, i.e., a shear from the x- and y- axis, using ecological stimuli. Immediate and long-term facilitation of fast adaptation induced by minutes time scales of extended skew exposures was tested. Fast adaptation was quantified via the magnitude of perceptual bias after a brief exposure to image skew in a constant stimulus procedure. Immediate facilitation was tested by comparing the magnitudes of fast adaptation that are measured on the same day before, i.e., baseline, and after extended skew exposure. The retention of the facilitation was evaluated by comparing the fast adaptation measured after, on average, 57 days of the previous extended skew exposure with the baseline. After one hour of skew exposure, the amount of fast adaptation significantly increased from the baseline measurement indicating immediate facilitation of the fast adaptation. This facilitation was retained at, on average, 57 days after the extended exposure. Thus, the results depicted experience dependent long-term facilitation of skew adaptation that potentially explains visual habituation to distortions of spectacles.

The ability to navigate through crowds of moving people accurately, efficiently, and without causing collisions is essential for our day-to-day lives. Vision provides key information about one's own self-motion as well as the motions of other people in the crowd. These two types of information (optic flow and biological motion) have each been investigated extensively; however, surprisingly little research has been dedicated to investigating how they are processed when presented concurrently. Here, we showed that patterns of biological motion have a negative impact on visual-heading estimation when people within the crowd move their limbs but do not move through the scene. Conversely, limb motion facilitates heading estimation when walkers move independently through the scene. Interestingly, this facilitation occurs for crowds containing both regular and perturbed depictions of humans, suggesting that it is likely caused by low-level motion cues inherent in the biological motion of other people.

When we move through the world, a pattern of expanding optic flow is generated on the retina. In completely rigid environments, this pattern signals one's direction of heading and is an important source of information for navigation. When we walk towards an oncoming person, the optic environment is not rigid, as the motion vectors generated by the other person represent a composite of that person's movement, his or her limb motion, and the observer's self-motion. Though this biological motion obfuscates the optic flow pattern, it also provides cues about the movement of other actors in the environment. It may be the case that the visual system takes advantage of these cues to simplify the decomposition of optic flow in the presence of other moving people. The current study sought to probe this possibility. In four experiments self-motion was simulated through an environment that was empty except for a single, walking point-light biological motion stimulus. We found that by using biological motion cues, observers were able to identify the presence of self-motion despite the lack of stable scene information. However, when estimating heading based on these stimuli, the pattern of observer heading estimates could be approximately reproduced by computing the vector sum of the walker's translation and the stimulated self-motion. This suggests that though biological motion can be used to disentangle self-motion in ambiguous situations, optic flow analysis does not use this information to derive heading estimates.

Rhythm and melody are two basic characteristics of music. Performing musicians have to pay attention to both, and avoid errors in either aspect of their performance. To investigate the neural processes involved in detecting melodic and rhythmic errors from auditory input we tested musicians on both kinds of deviations in a mismatch negativity (MMN) design. We found that MMN responses to a rhythmic deviation occurred at shorter latencies than MMN responses to a melodic deviation. Beamformer source analysis showed that the melodic deviation activated superior temporal, inferior frontal and superior frontal areas whereas the activation pattern of the rhythmic deviation focused more strongly on inferior and superior parietal areas, in addition to superior temporal cortex. Activation in the supplementary motor area occurred for both types of deviations. We also recorded responses to similar pitch and tempo deviations in a simple, non-musical repetitive tone pattern. In this case, there was no latency difference between the MMNs and cortical activation was smaller and mostly limited to auditory cortex. The results suggest that prediction and error detection of musical stimuli in trained musicians involve a broad cortical network and that rhythmic and melodic errors are processed in partially different cortical streams., (Copyright © 2015 Elsevier Inc. All rights reserved.)

Humans have a clear impression of facing in depth for point-light biological motion. However, this has not been measured systematically nor is it known on which cues humans rely for their judgment. In the present study subjects judged the facing orientation-in-depth of point-light displays. The displays represented natural walking and modified versions in which the time sequence was reversed, action was perturbed, the limbs and joints were nonrigid, the temporal sequence was scrambled, or the joint positions were scrambled. We found that the subjects were best at judging the facing direction of normal and reversed walking with an accuracy of 6° and 10° precision. The results show that pendular motion of the limb segments and the implicit knowledge of the human body play an important role for the precision of the judgment. Three further factors were relevant for the judgment of facing direction: (a) the discrimination of the front and back side, (b) the facing bias, and (c) the impression of depth from the display, probably due to the kinetic depth effect. The latter influences the accuracy, which differed strongly between subjects. The results suggest that the facing bias, to perceive the figure as facing toward the observer rather than away, is not related to the recognition of a human figure but rather to the presence of oscillating movements of the dots in the display.

Around the onset of a saccade toward a target, localization judgments are systematically biased toward the saccade endpoint. This perisaccadic compression is thought to be related to transsaccadic reorganization and due to interfering motor signals in visual maps. It has, however, only been investigated for saccades targeting a single target. Here, we examined whether saccade-sequence programming to stationary target(s) would affect the pattern of localization judgments of a briefly flashed stimulus. We presented saccade targets that could induce either a single saccade or two-saccade sequences and we flashed a bar around saccade onsets. For all two-saccade-sequence conditions, we showed that localization judgments of the stimulus flashed around the first saccade onset are biased toward an intermediate position between the two-saccade landing position, indicating the influence of the second saccade's parallel planning. This implies that motor-planning signals are most likely responsible for perisaccadic compression than motor execution feedback.

Perceptual phenomena that occur around the time of a saccade, such as peri-saccadic mislocalization or saccadic suppression of displacement, have often been linked to mechanisms of spatial stability. These phenomena are usually regarded as errors in processes of trans-saccadic spatial transformations and they provide important tools to study these processes. However, a true understanding of the underlying brain processes that participate in the preparation for a saccade and in the transfer of information across it requires a closer, more quantitative approach that links different perceptual phenomena with each other and with the functional requirements of ensuring spatial stability. We review a number of computational models of peri-saccadic spatial perception that provide steps in that direction. Although most models are concerned with only specific phenomena, some generalization and interconnection between them can be obtained from a comparison. Our analysis shows how different perceptual effects can coherently be brought together and linked back to neuronal mechanisms on the way to explaining vision across saccades.

In the temporal vicinity of a saccade onset, visual stability is transiently disrupted and briefly flashed visual stimuli undergo a systematic perceptual mislocalization. Specifically, when a stimulus is flashed around saccade onset, localization judgments are grossly biased toward the saccade endpoint. This peri-saccadic compression increases with saccade amplitude. Previous studies of peri-saccadic compression have typically used rather large saccade amplitudes. In the present study, we investigate systematically the pattern of errors for small saccade sizes (2°-10°), taking into account both the amplitude of the saccade and the position of the flashed stimulus (11 positions tested from 1 to 12°). Our results show a weaker compression effect for the smallest saccades. Moreover, we found that the strength of the compression depends on both stimulus side and relative distance to the saccade target.

The rapid and detailed recognition of human action from point-light displays is a remarkable ability and very robust against masking by motion signals. However, recognition of biological motion is strongly impaired when the typical point lights are replaced by pictures of complex objects. In a reaction time task and a detection in noise task, we asked subjects to decide if the walking direction is forward or backward. We found that complex objects as local elements impaired performance. When we compared different object categories, we found that human shapes as local objects gave more impairment than any other tested object category. Inverting or scrambling the human shapes restored the performance of walking perception. These results demonstrate an interference between object perception and biological motion recognition caused by shared processing capacities.

Walking through a crowd or driving on a busy street requires monitoring your own movement and that of others. The segmentation of these other, independently moving, objects is one of the most challenging tasks in vision as it requires fast and accurate computations for the disentangling of independent motion from egomotion, often in cluttered scenes. This is accomplished in our brain by the dorsal visual stream relying on heavy parallel-hierarchical processing across many areas. This study is the first to utilize the potential of such design in an artificial vision system. We emulate large parts of the dorsal stream in an abstract way and implement an architecture with six interdependent feature extraction stages (e.g., edges, stereo, optical flow, etc.). The computationally highly demanding combination of these features is used to reliably extract moving objects in real time. This way-utilizing the advantages of parallel-hierarchical design-we arrive at a novel and powerful artificial vision system that approaches richness, speed, and accuracy of visual processing in biological systems.

We demonstrate a strong sensory-motor coupling in visual localization in which experimental modification of the control of saccadic eye movements leads to an associated change in the perceived location of objects. Amplitudes of saccades to peripheral targets were altered by saccadic adaptation, induced by an artificial step of the saccade target during the eye movement, which leads the oculomotor system to recalibrate saccade parameters. Increasing saccade amplitudes induced concurrent shifts in perceived location of visual objects. The magnitude of perceptual shift depended on the size and persistence of errors between intended and actual saccade amplitudes. This tight agreement between the change of eye movement control and the change of localization shows that perceptual space is shaped by motor knowledge rather than simply constructed from visual input.

On winding roads, car drivers have to control speed and steering angle in order to keep the car in an optimal lane position. Among the strategies proposed for steering regulation are the use of the tangent point, a geometrical method, and gaze sampling, in which retinal flow lines obtained by tracking a spot on the future road need to be assessed. Previous studies used a variety of scenarios (real-road vs. simulator) and different road designs (closed vs. open bends, different curvatures) and found results speaking in favor of either strategy. Here, we investigate what effects the openness of the bend, i.e. the sight distance of the driver, has on the percentage with which drivers use the tangent point. Six drivers drove a test car repeatedly through a series of twelve bends on real roads while their eye-movements were recorded. Results show that the reliance on the tangent point is generally high and increases with the closedness (shorter sight distances) of the bend and higher curvature. In open bends they alternatively look far into the straight road segments adjacent to the bend, but do not use gaze sampling.

Studies with time-limited point-lights suggested that biological motion does not require local motion detection. These studies used walkers seen from the side, but biological motion perception excels also when walkers are oriented toward the observer, or in intermediate, half-profile views. In perspective projection, the local motion of points on the body provides a cue to the 3D structure of the walker. Thus, local point motion that was irrelevant for walkers in profile view may become important for biological motion perception in perspective projection. We compared performance on forward/backward walking discrimination of walkers in orthographic and perspective projection when view orientations and with point lifetime was varied. We found no difference between orthographic and perspective projections. Walkers with point lifetime 1 allowed forward-backward discrimination reliably in non-profile views, suggesting that local image motion is not required. Discrimination performance became extremely difficult in the frontal view, however. Follow-up experiments that tested lifetime, view orientation, and specific information from the feet indicated that this dependence on viewing angle can be explained by the reliance of the forward/backward discrimination on information about the movement of the lower legs, which is difficult to ascertain in the frontal view.

Biological motion, the movement of the human body presented by a small number of point lights, activates among other regions lining the posterior superior temporal sulcus (pSTS) and gyrus (pSTG) and of the fusiform gyrus. In previous studies with foveal stimuli the activity in the pSTS/pSTG was often confined to the right hemisphere and bilateral in fusiform gyrus. We presented biological motion stimuli in peripheral vision and measured the BOLD responses with functional MRI to test whether the right dominance in pSTS/pSTG also occurred with peripheral stimuli. We found activation exclusively in the right pSTG for both visual hemifields. In the fusiform gyrus activation was found in both hemispheres and for peripheral stimuli strongest for contralateral stimulation. However, in both fusiform gyri leftward-facing stimuli activated different subfields than rightward-facing stimuli, indicating a clustering of the selectivity for the orientation of the human body form. No such clustering was observed in the pSTG. The results indicate for the fusiform gyrus an organization with respect to the view orientation of the stimulus.

Car drivers can employ a number of strategies to negotiate curves. The tangent point strategy proposes the use of the angle between the tangent point of the inner lane markings and the car's current heading direction, which is proportional to the required steering angle. The gaze-sampling strategy proposes to fixate points on the future path and measure the curvature of optic flow vectors which can inform the drivers whether they over- or under-steer. Nine subjects drove repeatedly on the four loops of a motorway junction for which street parameters were available, while eye-movements, steering parameters and relations of the car to the lane were recorded. In the first half of the trials, we observed which strategy drivers normally use, whereas in the second half, we instructed subjects to use exclusively either the tangent-point or the gaze-sampling strategy and observed their steering behavior. Our results confirm that subjects normally look at the tangent point whereas they do not use gaze sampling of their own accord. Further, subjects drive more smoothly in terms of position on the lane and steering stability in the tangent-point condition.

Peri-saccadic perception experiments have revealed a multitude of mislocalization phenomena. For instance, a briefly flashed stimulus is perceived closer to the saccade target, whereas a displacement of the saccade target goes usually unnoticeable. This latter saccadic suppression of displacement has been explained by a built-in characteristic of the perceptual system: the assumption that during a saccade, the environment remains stable. We explored whether the mislocalization of a briefly flashed stimulus toward the saccade target also grounds in the built-in assumption of a stable environment. If the mislocalization of a peri-saccadically flashed stimulus originates from a post-saccadic alignment process, an additional location marker at the position of the upcoming flash should counteract compression. Alternatively, compression might be the result of peri-saccadic attentional phenomena. In this case, mislocalization should occur even if the position of the flashed stimulus is marked. When subjects were asked about their perceived location, they mislocalized the stimulus toward the saccade target, even though they were fully aware of the correct stimulus location. Thus, our results suggest that the uncertainty about the location of a flashed stimulus is not inherently relevant for compression.

Point-light biological motion stimuli provide spatio-temporal information about the structure of the human body in motion. Manipulation of the spatial structure of point-light stimuli reduces the ability of human observers to perceive biological motion. A recent study has reported that interference with the spatial structure of pointlight walkers also reduces the evoked eventrelated potentials over the occipitotemporal cortex, but that interference with the temporal structure of the stimuli evoked event-related potentials similar to normal biological motion stimuli. We systematically investigated the influence of spatial and temporal manipulation on 2 common discrimination tasks and compared it with predictions of a neurocomputational model previously proposed. This model first analyzes the spatial structure of the stimulus independently of the temporal information to derive body posture and subsequently analyzes the temporal sequence of body postures to derive movement direction. Similar to the model predictions, the psychophysical results show that human observers need only intact spatial configuration of the stimulus to discriminate the facing direction of a point-light walker. In contrast, movement direction discrimination needs a fully intact spatiotemporal pattern of the stimulus. The activation levels in the model predict the observed eventrelated potentials for the spatial and temporal manipulations.

Visual objects flashed before a saccade appear compressed toward the saccade target. Simultaneously flashed objects merge perceptually into one. To better understand cortical interactions in perisaccadic processing, we study the perception of features of mislocalized objects. We report four new findings: First, when multiple objects of different colors are compressed onto a single position, their color attributes remain distinguishable. Second, color attributes of objects compressed onto the same position compete for access to visual awareness. Third, objects presaccadically mislocalized onto a static background of identical color and luminance appear visible on top of that background. Object shape can be determined. Fourth, objects flashed during a saccade become invisible when a larger object is present at the mislocalized position. Thus, perisaccadic mislocalization affects the position of objects but retains other object features. Mislocalization must either occur in parallel to color and shape processing or at late stages of the visual pathway.