Time dilation effect in an active observer and virtual environment requires apparent motion: No dilation for retinal- or world-motion alone.

It is known that moving visual stimuli are perceived to last longer than stationary stimuli with the same physical duration (Kanai, Paffen, Hogendoorn, & Verstraten, 2006), and that motor actions (Tomassini & Morrone, 2016) and eye movements (Morrone, Ross, & Burr, 2005) can alter perceived duration. In the present work, we investigated the contributions of stimulus motion and self-motion to perceived duration while observers stood or walked in a virtual reality environment. Using a visual temporal reproduction task, we independently manipulated both the participants' motion (stationary or walking) and the stimulus motion (retinal stationary, real-world stationary and negative double velocity). When the observers were standing still, drifting gratings were perceived as lasting longer than duration-matched static gratings. Interestingly, we did not see any time distortion when observers were walking, neither when the gratings were kept stationary relative to the observer's point of view (i.e., no retinal motion) nor when they were stationary in the external world (i.e., producing the same retinal velocity as the walking condition with stationary grating). Self-motion caused significant dilation in perceived duration only when the gratings were moving at double speed, opposite to the observers' walking direction. Consistent with previous work (Fornaciai, Arrighi, & Burr, 2016), this suggests that the system is able to suppress self-generated motion to enhance external motion, which would have ecological benefits, for example, for threat detection while navigating through the environment.

The Lifetime of Salience Extends Beyond the Initial Saccade.

Several models of selection in search predict that saccades are biased toward conspicuous objects (also referred to as salient objects). Indeed, it has been demonstrated that initial saccades are biased toward the most conspicuous candidate. However, in a recent study, no such bias was found for the second saccade, and it was concluded that the attraction of conspicuous elements is limited to only short-latency initial saccades. This conclusion is based on only a single feature manipulation (orientation contrast) and conflicts with the prediction of influential salience models. Here, we investigate whether this result can be generalized beyond the domain of orientation. In displays containing three luminance annuli (Experiment 1), we find a considerable bias toward the most conspicuous candidate for the second saccade. In Experiment 1, the target could not be discriminated peripherally. When we made the target peripherally discriminable, the second saccade was no longer biased toward the more conspicuous candidate (Experiment 2). Thus, conspicuity plays a role in saccadic selection beyond the initial saccade. Whether second saccades are biased toward conspicuous objects appears to depend on the type of feature contrast underlying the conspicuity and the peripheral discriminability of target properties.

Intracranial Recordings of Occipital Cortex Responses to Illusory Visual Events.

UNLABELLED: Ambiguous visual stimuli elicit different perceptual interpretations over time, creating the illusion that a constant stimulus is changing. We investigate whether such spontaneous changes in visual perception involve occipital brain regions specialized for processing visual information, despite the absence of concomitant changes in stimulation. Spontaneous perceptual changes observed while viewing a binocular rivalry stimulus or an ambiguous structure-from-motion stimulus were compared with stimulus-induced perceptual changes that occurred in response to an actual stimulus change. Intracranial recordings from human occipital cortex revealed that spontaneous and stimulus-induced perceptual changes were both associated with an early transient increase in high-frequency power that was more spatially confined than a later transient decrease in low-frequency power. We suggest that the observed high-frequency and low-frequency modulations relate to initiation and maintenance of a percept, respectively. Our results are compatible with the idea that spontaneous changes in perception originate from competitive interactions within visual neural networks. SIGNIFICANCE STATEMENT: Ambiguous visual stimuli elicit different perceptual interpretations over time, creating the illusion that a constant stimulus is changing. The literature on the neural correlates of conscious visual perception remains inconclusive regarding the extent to which such spontaneous changes in perception involve sensory brain regions. In an attempt to bridge the gap between existing animal and human studies, we recorded from intracranial electrodes placed on the human occipital lobe. We compared two different kinds of ambiguous stimuli, binocular rivalry and the phenomenon of ambiguous structure-from-motion, enabling generalization of our findings across different stimuli. Our results indicate that spontaneous and stimulus-induced changes in perception (i.e., "illusory" and "real" changes in the stimulus, respectively) may involve sensory regions to a similar extent.

Revisiting the global effect and inhibition of return.

Saccades toward previously cued locations have longer latencies than saccades toward other locations, a phenomenon known as inhibition of return (IOR). Watanabe (Exp Brain Res 138:330-342. doi: 10.1007/s002210100709 , 2001) combined IOR with the global effect (where saccade landing points fall in between neighboring objects) to investigate whether IOR can also have a spatial component. When one of two neighboring targets was cued, there was a clear bias away from the cued location. In a condition where both targets were cued, it appeared that the global effect magnitude was similar to the condition without any cues. However, as the latencies in the double cue condition were shorter compared to the no cue condition, it is still an open question whether these results are representative for IOR. Considering the double cue condition can provide valuable insight into the interaction of the mechanisms underlying the two phenomena, here, we revisit this condition in an adapted paradigm. Our paradigm does result in longer latencies for the cued locations, and we find that the magnitude of the global effect is reduced significantly. Unexpectedly, this holds even when only including saccades with the same latencies for both conditions. Thus, the increased latencies associated with IOR cannot directly explain the reduction in global effect. The global effect reduction can likely best be seen as either a result of short-term depression of exogenous visual signals or a result of IOR established at the center of gravity of cues.

Interaction effects of visual stimulus speed and contrast on postural sway.

Manipulating the characteristics of visual stimuli that simulate self-motion through the environment can affect the resulting postural sway magnitude. In the present study, we address the question whether varying the contrast and speed of a linear translating dot pattern influences medial-lateral postural sway. In a first experiment, we investigated whether the postural sway magnitude increases with increasing dot speed, as was previously demonstrated for expanding and contracting stimuli. In a second experiment, we also manipulated the contrast of the stimuli. For reasons that high-contrast stimuli can be considered 'perceptually' stronger, we expect that higher-contrast stimuli induce more sway than lower-contrast stimuli. The results of the first experiment show that dot speed indeed influences postural sway, although in an unexpected way. For higher speeds, the sway is in the direction of the stimulus motion, yet for lower speeds the sway is in a direction opposite to the stimulus motion. The results of the second experiment show that dot contrast does affect postural sway, but that this depends on the speed of the moving dots. Interestingly, the direction of postural sway induced by a relatively low dot speed (4°/s) depends on dot contrast. Taken together, our results suggest that interactions between the visual, vestibular and proprioceptive system appear to be influenced by an internal representation of the visual stimulus, rather than being influenced by the external visual stimulus characteristics only.

Faces in Context: Does Face Perception Depend on the Orientation of the Visual Scene?

The mechanisms held responsible for familiar face recognition are thought to be orientation dependent; inverted faces are more difficult to recognize than their upright counterparts. Although this effect of inversion has been investigated extensively, researchers have typically sliced faces from photographs and presented them in isolation. As such, it is not known whether the perceived orientation of a face is inherited from the visual scene in which it appears. Here, we address this question by measuring performance in a simultaneous same-different task while manipulating both the orientation of the faces and the scene. We found that the face inversion effect survived scene inversion. Nonetheless, an improvement in performance when the scene was upside down suggests that sensitivity to identity increased when the faces were more easily segmented from the scene. Thus, while these data identify congruency with the visual environment as a contributing factor in recognition performance, they imply different mechanisms operate on upright and inverted faces.

Implicit perceptual memory modulates early visual processing of ambiguous images.

The way we perceive the present visual environment is influenced by past visual experiences. Here we investigated the neural basis of such experience dependency. We repeatedly presented human observers with an ambiguous visual stimulus (structure-from-motion) that can give rise to two distinct perceptual interpretations. Past visual experience is known to influence the perception of such stimuli. We recorded fast dynamics of neural activity shortly after stimulus onset using event-related electroencephalography. The number of previous occurrences of a certain percept modulated early posterior brain activity starting as early as 50 ms after stimulus onset. This modulation developed across hundreds of percept repetitions, reflecting several minutes of accumulating perceptual experience. Importantly, there was no such modulation when the mere number of previous stimulus presentations was considered regardless of how they were perceived. This indicates that the effect depended on previous perception rather than previous visual input. The short latency and posterior scalp location of the effect suggest that perceptual history modified bottom-up stimulus processing in early visual cortex. We propose that bottom-up neural responses to a given visual presentation are shaped, in part, by feedback modulation that occurred during previous presentations, thus allowing these responses to be biased in light of previous perceptual decisions.

Saccades toward the target are planned as sequences rather than as single steps.

To find a target during visual search, observers often need to make multiple eye movements, which results in a scan path. It is an open question whether the saccade destinations in scan paths are planned ahead. In the two experiments reported here, we investigated this question by focusing on the observer's ability to deviate from potentially planned paths. In the first experiment, the stimulus configuration could change during the initial saccade. We found that the observer's ability to deviate from potentially planned paths crucially depended on whether altered configurations could be processed with sufficient rapidity. In a follow-up experiment, we asked whether planned paths can include more than two saccade destinations. Investigating the influence of potentially planned paths on a secondary task demonstrated that planned paths can include at least three saccade destinations. Together, these experiments provide the first evidence of scan-path planning in visual search.

Walking modulates visual detection performance according to stride cycle phase.

Walking is among our most frequent and natural of voluntary behaviours, yet the consequences of locomotion upon perceptual and cognitive function remain largely unknown. Recent work has highlighted that although walking feels smooth and continuous, critical phases exist within each step for the successful coordination of perceptual and motor function. Here, we test whether these phasic demands impact upon visual perception, by assessing performance in a visual detection task during natural unencumbered walking. We finely sample visual performance over the stride cycle as participants walk along a smooth linear path at a comfortable speed in a wireless virtual reality environment. At the group-level, accuracy, reaction times, and response likelihood show strong oscillations, modulating at approximately 2 cycles per stride (~2 Hz) with a marked phase of optimal performance aligned with the swing phase of each step. At the participant level, Bayesian inference of population prevalence reveals highly prevalent oscillations in visual detection performance that cluster in two idiosyncratic frequency ranges (2 or 4 cycles per stride), with a strong phase alignment across participants.

Decoding the motion aftereffect in human visual cortex.

In the motion aftereffect (MAE), adapting to a moving stimulus causes a subsequently presented stationary stimulus to appear to move in the opposite direction. Recently, the neural basis of the motion aftereffect has received considerable interest, and a number of brain areas have been implicated in the generation of the illusory motion. Here, we use functional magnetic resonance imaging in combination with multivariate pattern classification to directly compare the neural activity evoked during the observation of both real and illusory motions. We show that the perceived illusory motion is not encoded in the same way as real motion in the same direction. Instead, suppression of the adapted direction of motion results in a shift of the population response of motion sensitive neurons in area MT+, resulting in activation patterns that are in fact more similar to real motion in orthogonal, rather than opposite directions. Although robust motion selectivity was observed in visual areas V1, V2, V3, and V4, this MAE-specific modulation of the population response was only observed in area MT+. Implications for our understanding of the motion aftereffect, and models of motion perception in general, are discussed.

Decreasing perceived optic flow rigidity increases postural sway.

Optic flow simulating self-motion through the environment can induce postural adjustments in observers. Some studies investigating this phenomenon have used optic flow patterns increasing in speed from center to periphery, whereas others used optic flow patterns with a constant speed. However, altering the speed gradient of an optic flow stimulus changes the perceived rigidity of such a stimulus. Optic flow stimuli that are perceived as rigid can be expected to provide a stronger sensation of self-motion than non-rigid optic flow, and this may well be reflected in the amount of postural sway. The current study, therefore, examined, by manipulating the speed gradient, to what extent the rigidity of an optic flow stimulus influences posture along the anterior-posterior axis. We used radial random dot expanding or contracting optic flow patterns with three different speed profiles (single-speed, linear speed gradient or quadratic speed gradient) that differentially induce the sensation of self-motion. Interestingly, most postural sway was observed for the non-rigid single-speed optic flow pattern, which contained the least self-motion information of the three profiles. Moreover, we found an anisotropy in that contracting optic flow produced more postural sway than expanding optic flow. In addition, the amount of postural sway increased with increasing stimulus speed, but for contracting optic flow only. Taken together, the results of the current study support the view that visual and sensorimotor systems appear to be tailored toward compensating for rigid optic flow stimulation.

How longer saccade latencies lead to a competition for salience.

It has been suggested that independent bottom-up and top-down processes govern saccadic selection. However, recent findings are hard to explain in such terms. We hypothesized that differences in visual-processing time can explain these findings, and we tested this using search displays containing two deviating elements, one requiring a short processing time and one requiring a long processing time. Following short saccade latencies, the deviation requiring less processing time was selected most frequently. This bias disappeared following long saccade latencies. Our results suggest that an element that attracts eye movements following short saccade latencies does so because it is the only element processed at that time. The temporal constraints of processing visual information therefore seem to be a determining factor in saccadic selection. Thus, relative saliency is a time-dependent phenomenon.

Saccadic selection and crowding in visual search: stronger lateral masking leads to shorter search times.

We investigated the role of crowding in saccadic selection during visual search. To guide eye movements, often information from the visual periphery is used. Crowding is known to deteriorate the quality of peripheral information. In four search experiments, we studied the role of crowding, by accompanying individual search elements by flankers. Varying the difference between target and flankers allowed us to manipulate crowding strength throughout the stimulus. We found that eye movements are biased toward areas with little crowding for conditions where a target could be discriminated peripherally. Interestingly, for conditions in which the target could not be discriminated peripherally, this bias reversed to areas with strong crowding. This led to shorter search times for a target presented in areas with stronger crowding, compared to a target presented in areas with less crowding. These findings suggest a dual role for crowding in visual search. The presence of flankers similar to the target deteriorates the quality of the peripheral target signal but can also attract eye movements, as more potential targets are present over the area.

Mapping the route to visual awareness.

The "neural correlate" of perceptual awareness is much sought-after. Here, we present an novel approach to the identification of possible neural correlates, in which we exploit the temporal connection that inevitably links the selection process that determines what we become aware of, and the development of awareness itself. Because the speed of selection determines when downstream processes can first become involved in generating awareness, the latency of neural processes provides a way to isolate the neural correlates of awareness. We recorded event-related potentials (ERPs) while observers carried out a visual behavioral task designed to estimate attentional selection latency. We show that within-task trial-by-trial behavioral variability in attentional selection latency correlates to trial-by-trial variability in ERP latency. This was true in a posterior contralateral region, and in central and frontal areas, thereby implicating these as waypoints along which visual information flows on the way to visual awareness.

Perceptual benefits of objecthood.

Object-based attention facilitates the processing of features that form the object. Two hypotheses are conceivable for how object-based attention is deployed to an object's features: first, the object is attended by selecting its features; alternatively, a configuration of features as such is attended by selecting the object representation they form. Only for the latter alternative, the perception of a feature configuration as entity ("objecthood") is a necessary condition for object-based attention. Disentangling the two alternatives requires the comparison of identical feature configurations that induce the perception of an object in one condition ("bound") and do not do so in another condition ("unbound"). We used an ambiguous stimulus, whose percept spontaneously switches between bound and unbound, while the stimulus itself remains unchanged. We tested discrimination on the boundary of the diamond as well as detection of probes inside and outside the diamond. We found discrimination performance to be increased if features were perceptually bound into an object. Furthermore, detection performance was higher within and lower outside the bound object as compared to the unbound configuration. Consequently, the facilitation of processing by object-based attention requires objecthood, that is, a unified internal representation of an "object"-not a mere collection of features.

Suppressed images selectively affect the dominant percept during binocular rivalry.

During binocular rivalry, perception alternates between dissimilar images that are presented dichoptically. It has been argued that perception during the dominance phase of rivalry is unaffected by the suppressed image. Recent evidence suggests, however, that the suppressed image does affect perception of the dominant image, yet the extent and nature of this interaction remain elusive. We hypothesize that this interaction depends on the difference in feature content between the rivaling images. Here, we investigate how sensitivity to probes presented in the image that is currently dominant in perception is affected by the suppressed image. Observers performed a 2AFC discrimination task on oriented probes (Experiment 1) or probes with different motion directions (Experiment 2). Our results show that performance on both orientation and motion direction discrimination was affected by the content of the suppressed image. The strength of interference depended specifically on the difference in feature content (e.g., the difference in orientation) between the probe and the suppressed image. Moreover, the pattern of interference by the suppressed image is qualitatively similar to the situation where this image and the probe are simultaneously visible. We conclude that perception during the dominance phase of rivalry is affected by a suppressed image as if it were visible.

Vestibular and active self-motion signals drive visual perception in binocular rivalry.

Multisensory integration helps the brain build reliable models of the world and resolve ambiguities. Visual interactions with sound and touch are well established but vestibular influences on vision are less well studied. Here, we test the vestibular influence on vision using horizontally opposed motions presented one to each eye so that visual perception is unstable and alternates irregularly. Passive, whole-body rotations in the yaw plane stabilized visual alternations, with perceived direction oscillating congruently with rotation (leftward motion during leftward rotation, and vice versa). This demonstrates a purely vestibular signal can resolve ambiguous visual motion and determine visual perception. Active self-rotation following the same sinusoidal profile also entrained vision to the rotation cycle - more strongly and with a lesser time lag, likely because of efference copy and predictive internal models. Both experiments show that visual ambiguity provides an effective paradigm to reveal how vestibular and motor inputs can shape visual perception.