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Our visual system faces the challenging task to construct integrated visual representations from the visual input projected on our retinae. Previous research has provided mixed evidence as to whether visual awareness of the stimulus parts is required for such integration to occur. Here, we address this issue by taking a novel approach in which we combine a monocular rivalry stimulus (i.e., a bistable rotating cylinder) with binocular rivalry. The results of Experiment 1 show that in a rivalry condition, where one half of the cylinder is perceptually suppressed, significantly more perceptual switches occur that are consistent with visual integration of the whole cylinder than occur in a control condition, where only half of the cylinder is presented at a time and the presentation of the two images is physically alternated. In Experiment 2, stimulation in the observer's dominant eye was kept dominant by presenting the half cylinder in this eye at higher contrast and by surrounding it with a flickering context. Results show that the strong convexity bias that was found in a control condition, where no stimulus was presented in the suppressed eye, almost completely disappears when the unseen half is presented in the suppressed eye, indicating that both halves visually integrate and, subsequently, compete for convexity. These findings provide evidence that unseen visual information is biased towards a representation that is congruent with the current visible representation and, hence, that principles of perceptual organization also apply to parts of the visual input that remain unseen by the observer.

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., (Copyright © 2016 the authors 0270-6474/16/366297-15$15.00/0.)

The plasticity of the human brain, as shown in perceptual learning, is generally reflected by improved task performance after training. Here, we show that perceptual suppression can be increased through training. In the first experiment, binocular rivalry suppression of a specific orientation was trained, leading to a relative reduction in sensitivity to the trained orientation. In a second experiment, two orthogonal orientations were suppressed in alternating training blocks, in the left and right eye, respectively. This double-training procedure lead to reduced sensitivity for the orientation that was suppression-trained in each specific eye, implying that training of feature suppression is specific for the eye in which the oriented grating was presented during training. Results of a control experiment indicate that the obtained effects are indeed due to suppression during training, instead of being merely due to the repetitive presentation of the oriented gratings. Visual plasticity is essential for a person's visual development. The finding that plasticity can result in increased perceptual suppression reported here may prove to be significant in understanding human visual development. It emphasizes that for stable vision, not only the enhancement of relevant signals is crucial, but also the reliable and stable suppression of (task) irrelevant signals.

The research presented here focuses on the influence of background music on the beer-tasting experience. An experiment is reported in which different groups of customers tasted a beer under three different conditions (N = 231). The control group was presented with an unlabeled beer, the second group with a labeled beer, and the third group with a labeled beer together with a customized sonic cue (a short clip from an existing song). In general, the beer-tasting experience was rated as more enjoyable with music than when the tasting was conducted in silence. In particular, those who were familiar with the band that had composed the song, liked the beer more after having tasted it while listening to the song, than those who knew the band, but only saw the label while tasting. These results support the idea that customized sound-tasting experiences can complement the process of developing novel beverage (and presumably also food) events. We suggest that involving musicians and researchers alongside brewers in the process of beer development, offers an interesting model for future development. Finally, we discuss the role of attention in sound-tasting experiences, and the importance that a positive hedonic reaction toward a song can have for the ensuing tasting experience.

Does one need to be aware of a visual stimulus for it to be perceptually organized into a coherent whole? The answer to this question regarding the interplay between Gestalts and visual awareness remains unclear. Using interocular suppression as the paradigm for rendering stimuli invisible, conflicting evidence has been obtained as to whether the traditional Kanizsa surface is constructed during interocular suppression. While Sobel and Blake (2003) and Harris, Schwarzkopf, Song, Bahrami, and Rees (2011) failed to find evidence for this, Wang, Weng, and He (2012) showed that standard configurations of Kanizsa pacmen would break interocular suppression faster than their rotated counterparts. In the current study, we replicated the findings by Wang et al. (2012) but show that neither an account based on the construction of a surface nor one based on the long-range collinearities in the standard Kanizsa configuration stimulus could fully explain the difference in breakthrough times. We discuss these findings in the context of differences in the amplitudes of the Fourier orientation spectra for all stimulus types. Thus, we find no evidence that the integration of separate elements takes place during interocular suppression of Kanizsa stimuli, suggesting that this Gestalt involving figure-ground assignment is not constructed when rendered nonconscious using interocular suppression.

The aim of this systematic review was to integrate and assess evidence for the effectiveness of multisensory stimulation (i.e., stimulating at least two of the following sensory systems: visual, auditory, and somatosensory) as a possible rehabilitation method after stroke. Evidence was considered with a focus on low-level, perceptual (visual, auditory and somatosensory deficits), as well as higher-level, cognitive, sensory deficits. We referred to the electronic databases Scopus and PubMed to search for articles that were published before May 2015. Studies were included which evaluated the effects of multisensory stimulation on patients with low- or higher-level sensory deficits caused by stroke. Twenty-one studies were included in this review and the quality of these studies was assessed (based on eight elements: randomization, inclusion of control patient group, blinding of participants, blinding of researchers, follow-up, group size, reporting effect sizes, and reporting time post-stroke). Twenty of the twenty-one included studies demonstrate beneficial effects on low- and/or higher-level sensory deficits after stroke. Notwithstanding these beneficial effects, the quality of the studies is insufficient for valid conclusion that multisensory stimulation can be successfully applied as an effective intervention. A valuable and necessary next step would be to set up well-designed randomized controlled trials to examine the effectiveness of multisensory stimulation as an intervention for low- and/or higher-level sensory deficits after stroke. Finally, we consider the potential mechanisms of multisensory stimulation for rehabilitation to guide this future research.

Research on visual rivalry has demonstrated that consecutive dominance durations are serially dependent, implying that the underlying competition mechanism is not driven by some random process but includes a memory component. Here we asked whether serial dependence is also observed in continuous flash suppression (CFS). We addressed this question by analyzing a large dataset of time series of suppression durations obtained in a series of so-called "breaking CFS" experiments in which the duration of the period is measured until a suppressed target breaks through the CFS mask. Across experimental manipulations, stimuli, and observers, we found that (i) the distribution of breakthrough rates was fit less well by a gamma distribution than in conventional visual rivalry paradigms, (ii) the suppression duration on a previous trial influenced the suppression duration on a later trial up to as long as a lag of eight trials, and (iii) the mechanism underlying these serial correlations was predominantly monocular. We conclude that the underlying competition mechanism of CFS also includes a memory component that is primarily, but not necessarily exclusively, monocular in nature. We suggest that the temporal dependency structure of suppression durations in CFS is akin to those observed in binocular rivalry, which might imply that both phenomena tap into similar rather than distinct mechanisms.

The soundscapes of those places where we eat and drink can influence our perception of taste. Here, we investigated whether contextual sound would enhance the subjective value of a tasting experience. The customers in a chocolate shop were invited to take part in an experiment in which they had to evaluate a chocolate's taste while listening to an auditory stimulus. Four different conditions were presented in a between-participants design. Envisioning a more ecological approach, a pre-recorded piece of popular music and the shop's own soundscape were used as the sonic stimuli. The results revealed that not only did the customers report having a significantly better tasting experience when the sounds were presented as part of the food's identity, but they were also willing to pay significantly more for the experience. The method outlined here paves a new approach to dealing with the design of multisensory tasting experiences, and gastronomic situations.

Synthetic amorphous silica (SAS) like NM-200 is used in a wide variety of technological applications and consumer products. Although SAS has been widely investigated the available reproductive toxicity studies are old and do not cover all requirements of current OECD Guidelines. As part of a CEFIC-LRI project, NM-200 was tested in a two-generation reproduction toxicity study according to OECD guideline 416. Male and female rats were treated by oral gavage with NM-200 at dose levels of 0, 100, 300 and 1000mg/kg bw/day for two generations. Body weight and food consumption were measured throughout the study. Reproductive and developmental parameters were measured and at sacrifice (reproductive) organs and tissues were sampled for histopathological analysis. Oral administration of NM-200 up to 1000mg/kg bw/day had no adverse effects on the reproductive performance of rats or on the growth and development of the offspring into adulthood for two consecutive generations. The NOAEL was 1000mg/kg body weight per day., (Copyright © 2015 Elsevier Inc. All rights reserved.)

Previous studies using binocular rivalry have shown that signals in a modality other than the visual can bias dominance durations depending on their congruency with the rivaling stimuli. More recently, studies using continuous flash suppression (CFS) have reported that multisensory integration influences how long visual stimuli remain suppressed. In this study, using CFS, we examined whether the contrast thresholds for detecting visual looming stimuli are influenced by a congruent auditory stimulus. In Experiment 1, we show that a looming visual stimulus can result in lower detection thresholds compared to a static concentric grating, but that auditory tone pips congruent with the looming stimulus did not lower suppression thresholds any further. In Experiments 2, 3, and 4, we again observed no advantage for congruent multisensory stimuli. These results add to our understanding of the conditions under which multisensory integration is possible, and suggest that certain forms of multisensory integration are not evident when the visual stimulus is suppressed from awareness using CFS.

Segmentation of a visual scene in "figure" and "ground" is essential for perception of the three-dimensional layout of a scene. In cases of bi-stable perception, two distinct figure-ground interpretations alternate over time. We were interested in the temporal dynamics of these alternations, in particular when the same image is presented repeatedly, with short blank periods in-between. Surprisingly, we found that the intermittent presentation of Rubin's classical "face-or-vase" figure, which is frequently taken as a standard case of bi-stable figure-ground perception, often evoked perceptual switches during the short presentations and stabilization was not prominent. Interestingly, bi-stable perception of Kanizsa's anomalous transparency figure did strongly stabilize across blanks. We also found stabilization for the Necker cube, which we used for comparison. The degree of stabilization (and the lack of it) varied across stimuli and across individuals. Our results indicate, against common expectation, that the stabilization phenomenon cannot be generally evoked by intermittent presentation. We argue that top-down feedback factors such as familiarity, semantics, expectation, and perceptual bias contribute to the complex processes underlying the temporal dynamics of bi-stable figure-ground perception., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

After hearing a tone, the human auditory system becomes more sensitive to similar tones than to other tones. Current auditory models explain this phenomenon by a simple bandpass attention filter. Here, we demonstrate that auditory attention involves multiple pass-bands around octave-related frequencies above and below the cued tone. Intriguingly, this "octave effect" not only occurs for physically presented tones, but even persists for the missing fundamental in complex tones, and for imagined tones. Our results suggest neural interactions combining octave-related frequencies, likely located in nonprimary cortical regions. We speculate that this connectivity scheme evolved from exposure to natural vibrations containing octave-related spectral peaks, e.g., as produced by vocal cords.

Emotionally arousing stimuli are perceived and remembered better than neutral stimuli. Under threat, this negativity bias is further increased. We investigated whether working memory (WM) load can attenuate incidental memory for emotional images. Two groups of participants performed the N-back task with two WM load levels. In one group, we induced anxiety using a threat of shock paradigm to increase attentional processing of negative information. During task performance we incidentally and briefly flashed emotional distracter images which prolonged response times in both load conditions. A subsequent unannounced immediate recognition memory test revealed that when load at exposure had been low, recognition was better for negative items in both participant groups. This enhancement, however, was attenuated under high load, leaving performance on neutral items unchanged regardless of the threat of shock manipulation. We conclude that both in threat and in normal states WM load at exposure can attenuate immediate emotional memory enhancement.

Successful interactions with the environment entail interpreting ambiguous sensory information. To address this challenge it has been suggested that the brain optimizes performance through experience. Here we used functional magnetic resonance imaging (fMRI) to investigate whether perceptual experience modulates the cortical circuits involved in visual awareness. Using ambiguous visual stimuli (binocular rivalry or ambiguous structure-from-motion) we were able to disentangle the co-occurring influences of stimulus repetition and perceptual repetition. For both types of ambiguous stimuli we observed that the mere repetition of the stimulus evoked an entirely different pattern of activity modulations than the repetition of a particular perceptual interpretation of the stimulus. Regarding stimulus repetition, decreased fMRI responses were evident during binocular rivalry but weaker during 3-D motion rivalry. Perceptual repetition, on the other hand, entailed increased activity in stimulus-specific visual brain regions - for binocular rivalry in the early visual regions and for ambiguous structure-from-motion in both early as well as higher visual regions. This indicates that the repeated activation of a visual network mediating a particular percept facilitated its later reactivation. Perceptual repetition was also associated with a response change in the parietal cortex that was similar for the two types of ambiguous stimuli, possibly relating to the temporal integration of perceptual information. We suggest that perceptual repetition is associated with a facilitation of neural activity within and between percept-specific visual networks and parietal networks involved in the temporal integration of perceptual information, thereby enhancing the stability of previously experienced percepts., (© 2012 Federation of European Neuroscience Societies and Blackwell Publishing Ltd.)

Ambiguous visual stimuli provide the brain with sensory information that contains conflicting evidence for multiple mutually exclusive interpretations. Two distinct aspects of the phenomenological experience associated with viewing ambiguous visual stimuli are the apparent stability of perception whenever one perceptual interpretation is dominant, and the instability of perception that causes perceptual dominance to alternate between perceptual interpretations upon extended viewing. This review summarizes several ways in which contextual information can help the brain resolve visual ambiguities and construct temporarily stable perceptual experiences. Temporal context through prior stimulation or internal brain states brought about by feedback from higher cortical processing levels may alter the response characteristics of specific neurons involved in rivalry resolution. Furthermore, spatial or crossmodal context may strengthen the neuronal representation of one of the possible perceptual interpretations and consequently bias the rivalry process towards it. We suggest that contextual influences on perceptual choices with ambiguous visual stimuli can be highly informative about the neuronal mechanisms of context-driven inference in the general processes of perceptual decision-making.

Observers continually make unconscious inferences about the state of the world based on ambiguous sensory information. This process of perceptual decision-making may be optimized by learning from experience. We investigated the influence of previous perceptual experience on the interpretation of ambiguous visual information. Observers were pre-exposed to a perceptually stabilized sequence of an ambiguous structure-from-motion stimulus by means of intermittent presentation. At the subsequent re-appearance of the same ambiguous stimulus perception was initially biased toward the previously stabilized perceptual interpretation. However, prolonged viewing revealed a bias toward the alternative perceptual interpretation. The prevalence of the alternative percept during ongoing viewing was largely due to increased durations of this percept, as there was no reliable decrease in the durations of the pre-exposed percept. Moreover, the duration of the alternative percept was modulated by the specific characteristics of the pre-exposure, whereas the durations of the pre-exposed percept were not. The increase in duration of the alternative percept was larger when the pre-exposure had lasted longer and was larger after ambiguous pre-exposure than after unambiguous pre-exposure. Using a binocular rivalry stimulus we found analogous perceptual biases, while pre-exposure did not affect eye-bias. We conclude that previously perceived interpretations dominate at the onset of ambiguous sensory information, whereas alternative interpretations dominate prolonged viewing. Thus, at first instance ambiguous information seems to be judged using familiar percepts, while re-evaluation later on allows for alternative interpretations.

In bistable vision, one constant ambiguous stimulus leads to 2 alternating conscious percepts. This perceptual switching occurs spontaneously but can also be influenced through voluntary control. Neuroimaging studies have reported that frontal regions are activated during spontaneous perceptual switches, leading some researchers to suggest that frontal regions causally induce perceptual switches. But the opposite also seems possible: frontal activations may themselves be caused by spontaneous switches. Classically implicated in attentional processes, these same regions are also candidates for the origins of voluntary control over bistable vision. Here too, it remains unknown whether frontal cortex is actually functionally relevant. It is even possible that spontaneous perceptual switches and voluntarily induced switches are mediated by the same top-down mechanisms. To directly address these issues, we here induced "virtual lesions," with transcranial magnetic stimulation, in frontal, parietal, and 2 lower level visual cortices using an established ambiguous structure-from-motion stimulus. We found that dorsolateral prefrontal cortex was causally relevant for voluntary control over perceptual switches. In contrast, we failed to find any evidence for an active role of frontal cortex in passive bistable vision. Thus, it seems the same pathway used for willed top-down modulation of bistable vision is not used during passive bistable viewing.

When sensory input allows for multiple, competing perceptual interpretations, observers' perception can fluctuate over time, which is called bistable perception. Imaging studies in humans have revealed transient responses in a right-lateralized network in the frontal-parietal cortex (rFPC) around the time of perceptual transitions between interpretations, potentially reflecting the neural initiation of transitions. We investigated the role of this activity in male human observers, with specific interest in its relation to the temporal structure of transitions, which can be either instantaneous or prolonged by periods during which observers experience a mix of both perceptual interpretations. Using both bistable apparent motion and binocular rivalry, we show that transition-related rFPC activity is larger for transitions that last longer, suggesting that rFPC remains active as long as a transition lasts. We also replicate earlier findings that rFPC activity during binocular rivalry transitions exceeds activity during yoked transitions that are simulated using video replay. However, we show that this established finding holds only when perceptual transitions are replayed as instantaneous events. When replay, instead, depicts transitions with the actual durations reported during rivalry, yoked transitions and genuine rivalry transitions elicit equal activity. Together, our results are consistent with the view that at least a component of rFPC activation during bistable perception reflects a response to perceptual transitions, both real and yoked, rather than their cause. This component of activity could reflect the change in sensory experience and task demand that occurs during transitions, which fits well with the known role of these areas in attention and decision making.

Stereo-vision is generally considered to provide information about depth in a visual scene derived from disparities in the positions of an image on the two eyes; a new study has found evidence that retinal-image coding relative to the head is also important., (Copyright 2010 Elsevier Ltd. All rights reserved.)

A previous study has shown that diverting attention from binocular rivalry to a visual distractor task results in a slowing of rivalry alternation rate between simple orthogonal orientations. Here, we investigate whether the slowing of visual perceptual alternations will occur when attention is diverted to an auditory distractor task, and we extend the investigation by testing this for two kinds of binocular rivalry stimuli and for the Necker cube. Our results show that doing the auditory attention task does indeed slow visual perceptual alternations, that the slowing effect is a graded function of attentional load, and that the attentional slowing effect is less pronounced for grating rivalry than for house/face rivalry and for the Necker cube. These results are explained in terms of supramodal attentional resources modulating a high-level interpretative process in perceptual ambiguity, together with a role for feedback to early visual processes in the case of binocular rivalry., (Copyright 2010 Elsevier Ltd. All rights reserved.)

The issue of the existence of planes-understood as the carriers of a nexus of straight lines-in the monocular visual space of a stationary human observer has never been addressed. The most recent empirical data apply to binocular visual space and date from the 1960s (Foley, 1964). This appears to be both the first and the last time this basic issue was addressed empirically. Yet the question is of considerable conceptual interest. Here we report on a direct empirical test of the existence of planes in monocular visual space for a group of sixteen experienced observers. For the majority of these observers monocular visual space lacks a projective structure, albeit in qualitatively different ways. This greatly reduces the set of viable geometrical models. For example, it rules out all the classical homogeneous spaces (the Cayley-Klein geometries) such as the familiar Luneburg model. The qualitatively different behavior of experienced observers implies that the generic population might well be inhomogeneous with respect to the structure of visual space., (Copyright 2009 Elsevier B.V. All rights reserved.)

When faced with inconclusive or conflicting visual input human observers experience one of multiple possible perceptions. One factor that determines perception of such an ambiguous stimulus is how the same stimulus was perceived on previous occasions, a phenomenon called perceptual memory. We examined perceptual memory of an ambiguous motion stimulus while applying transcranial magnetic stimulation (TMS) to the motion-sensitive areas of the middle temporal cortex (hMT+). TMS increased the predominance of whichever perceptual interpretation was most commonly reported by a given observer at baseline, with reduced perception of the less favored interpretation. This increased incidence of the preferred percept indicates impaired long-term buildup of perceptual memory traces that normally act against individual percept biases. We observed no effect on short-term memory traces acting from one presentation to the next. Our results indicate that hMT+ is important for the long-term buildup of perceptual memory for ambiguous motion stimuli.

When the sensory system is subjected to ambiguous input, perception alternates between interpretations in a seemingly random fashion. Although neuronal noise obviously plays a role, the neural mechanism for the generation of randomness at the slow time scale of the percept durations (multiple seconds) is unresolved. Here significant nonzero serial correlations are reported in series of visual percept durations (to the author's knowledge for the first time accounting for duration impurities caused by reaction time, drift, and incomplete percepts). Serial correlations for perceptual rivalry using structure-from-motion ambiguity were smaller than for binocular rivalry using orthogonal gratings. A spectrum of computational models is considered, and it is concluded that noise in adaptation of percept-related neurons causes the serial correlations. This work bridges, in a physiologically plausible way, widely appreciated deterministic modeling and randomness in experimental observations of visual rivalry.

The neural mechanisms underlying attentional selection of competing neural signals for awareness remains an unresolved issue. We studied attentional selection, using perceptually ambiguous stimuli in a novel multisensory paradigm that combined competing auditory and competing visual stimuli. We demonstrate that the ability to select, and attentively hold, one of the competing alternatives in either sensory modality is greatly enhanced when there is a matching cross-modal stimulus. Intriguingly, this multimodal enhancement of attentional selection seems to require a conscious act of attention, as passively experiencing the multisensory stimuli did not enhance control over the stimulus. We also demonstrate that congruent auditory or tactile information, and combined auditory-tactile information, aids attentional control over competing visual stimuli and visa versa. Our data suggest a functional role for recently found neurons that combine voluntarily initiated attentional functions across sensory modalities. We argue that these units provide a mechanism for structuring multisensory inputs that are then used to selectively modulate early (unimodal) cortical processing, boosting the gain of task-relevant features for willful control over perceptual awareness.

When observing bistable stimuli, the percept can change in the absence of changes in the stimulus itself. When intermittently presenting a bistable stimulus, the number of perceptual alternations can increase or decrease, depending on the duration of the period that the stimulus is removed from screen between stimulus presentations (off-period). Longer off-periods lead to stabilization of the percept, while short off-periods produce perceptual alternations. Here we compare fMRI brain activation across percept repetitions and alternations when observing an intermittently presented ambiguously rotating structure from motion sphere. In the first experimental session, subjects were requested to voluntarily control the percept into either a repeating or an alternating perceptual regime at a single off-period. In a consecutive session, subjects observed the sphere uninstructed, and reported alternations and repetitions. The behavioral data showed that there were marked individual biases for observing the sphere as either repeating or alternating. The fMRI data showed activation differences between alternating and repeating perceptual regimes in an extensive network that included parietal cortex, dorsal premotor area, dorsolateral prefrontal cortex, supplementary motor area, insula, and cerebellum. However, these activation differences could all be explained by intersubject differences in the bias for one of the two perceptual regimes. The stronger the bias was for a particular perceptual regime, the less activation and vice versa. We conclude that widespread activation differences between perceptual regimes can be accounted for by differences in the perceptual bias for one of the two regimes.

We employed a parametric psychophysical design in combination with functional imaging to examine the influence of metric changes in perceptual incongruence on perceptual alternation rates and cortical responses. Subjects viewed a bistable stimulus defined by incongruent depth cues; bistability resulted from incongruence between binocular disparity and monocular perspective cues that specify different slants (slant rivalry). Psychophysical results revealed that perceptual alternation rates were positively correlated with the degree of perceived incongruence. Functional imaging revealed systematic increases in activity that paralleled the psychophysical results within anterior intraparietal sulcus, prior to the onset of perceptual alternations. We suggest that this cortical activity predicts the frequency of subsequent alternations, implying a putative causal role for these areas in initiating bistable perception. In contrast, areas implicated in form and depth processing (LOC and V3A) were sensitive to the degree of slant, but failed to show increases in activity when these cues were in conflict.

Adaptation is one of the key constituents of the perceptual alternation process during binocular rivalry, as it has been shown that preadapting one of the rivaling pairs before rivalry onset biases perception away from the adapted stimulus during rivalry. We investigated the influence of retinotopic and spatiotopic preadaptation on binocular rivalry. We show that for grating stimuli, preadaptation only influences rivalry when adaptation and rivalry locations are retinotopically matched. With more complex house and face stimuli, effects of preadaptation are found for both retinotopic and spatiotopic preadaptation, showing the importance of spatiotopic encoding in binocular rivalry. We show, furthermore, that adaptation to phase-scrambled faces results in retinotopic effects only, demonstrating the importance of form content for spatiotopic adaptation effects, as opposed to spatial frequency content. Are the spatiotopic adaptation influences on rivalry caused by direct spatiotopic stimulus interactions, or instead are they due to altered feedback from the adapted spatiotopic representations to the retinotopic representations that are involved in rivalry? By using rivaling face and grating stimuli that minimize rivalry between spatiotopic representations while still engaging these representations in stimulus encoding, we show that at least part of the preadaptation effects with face stimuli depend on feedback information.

At the onset of bistable stimuli, the brain needs to choose which of the competing perceptual interpretations will first reach awareness. Stimulus manipulations and cognitive control both influence this choice process, but the underlying mechanisms and interactions remain poorly understood. Using intermittent presentation of bistable visual stimuli, we demonstrate that short interruptions cause perceptual reversals upon the next presentation, whereas longer interstimulus intervals stabilize the percept. Top-down voluntary control biases this process but does not override the timing dependencies. Extending a recently introduced low-level neural model, we demonstrate that percept-choice dynamics in bistable vision can be fully understood with interactions in early neural processing stages. Our model includes adaptive neural processing preceding a rivalry resolution stage with cross-inhibition, adaptation, and an interaction of the adaptation levels with a neural baseline. Most importantly, our findings suggest that top-down attentional control over bistable stimuli interacts with low-level mechanisms at early levels of sensory processing before perceptual conflicts are resolved and perceptual choices about bistable stimuli are made.

We investigated the spatio-temporal dynamics of the steady-state-visual-evoked field during perceptual switches in binocular rivalry using MEG. Several authors have previously used frequency tagging in MEG studies on binocular rivalry and have claimed to have found a widespread network of synchronized areas that are entrained by the stimulus, reaching up to frontal regions. Moreover, it has been claimed that the results prove that becoming aware of a stimulus is established by increased intra- and interhemispheric synchronization of brain areas, separated by large distances. Our results dovetailed nicely with previous findings such as power and coherence modulations as a function of perceptual state. However, while we also found stimulus-entrained activity across the entire scalp, a phase analysis revealed that the spatially extended nature of the frequency tag was produced by a limited set of occipital sources. Furthermore, we provide evidence that the coherence results from earlier studies are ambiguous in that they likely measured coherence between different sensors whose signals were dominated by the same sources. We conclude that the claims about widespread synchronized networks to consciously perceive flicker stimuli are currently unconvincing.

When visual input is inconclusive, does previous experience aid the visual system in attaining an accurate perceptual interpretation? Prolonged viewing of a visually ambiguous stimulus causes perception to alternate between conflicting interpretations. When viewed intermittently, however, ambiguous stimuli tend to evoke the same percept on many consecutive presentations. This perceptual stabilization has been suggested to reflect persistence of the most recent percept throughout the blank that separates two presentations. Here we show that the memory trace that causes stabilization reflects not just the latest percept, but perception during a much longer period. That is, the choice between competing percepts at stimulus reappearance is determined by an elaborate history of prior perception. Specifically, we demonstrate a seconds-long influence of the latest percept, as well as a more persistent influence based on the relative proportion of dominance during a preceding period of at least one minute. In case short-term perceptual history and long-term perceptual history are opposed (because perception has recently switched after prolonged stabilization), the long-term influence recovers after the effect of the latest percept has worn off, indicating independence between time scales. We accommodate these results by adding two positive adaptation terms, one with a short time constant and one with a long time constant, to a standard model of perceptual switching.

Presenting the eyes with spatially mismatched images causes a phenomenon known as binocular rivalry-a fluctuation of awareness whereby each eye's image alternately determines perception. Binocular rivalry is used to study interocular conflict resolution and the formation of conscious awareness from retinal images. Although the spatial determinants of rivalry have been well-characterized, the temporal determinants are still largely unstudied. We confirm a previous observation that conflicting images do not need to be presented continuously or simultaneously to elicit binocular rivalry. This process has a temporal limit of about 350 ms, which is an order of magnitude larger than the visual system's temporal resolution. We characterize this temporal limit of binocular rivalry by showing that it is independent of low-level information such as interocular timing differences, contrast-reversals, stimulus energy, and eye-of-origin information. This suggests the temporal factors maintaining rivalry relate more to higher-level form information, than to low-level visual information. Systematically comparing the role of form and motion-the processing of which may be assigned to ventral and dorsal visual pathways, respectively-reveals that this temporal limit is determined by form conflict rather than motion conflict. Together, our findings demonstrate that binocular conflict resolution depends on temporally coarse form-based processing, possibly originating in the ventral visual pathway.

We studied the influence of perceived surface orientation on vergence accompanying a saccade while viewing an ambiguous stimulus. We used the slant rivalry stimulus, in which perspective foreshortening and disparity specified opposite surface orientations. This rivalrous configuration induces alternations of perceived surface orientation, while the slant cues remain constant. Subjects were able to voluntarily control their perceptual state while viewing the ambiguous stimulus. They were asked to make a saccade across the perceived slanted surface. Our data show that vergence responses closely approximated the vergence response predicted by the disparity cue, irrespective of voluntarily controlled perceived orientation. However, comparing the data obtained while viewing the ambiguous stimulus with data from an unambiguous stimulus condition (when disparity and perspective specified similar surface orientations) revealed an effect of perspective cues on vergence. Collectively our results show that depth cues rather than perceived depth govern vergence.

The mechanisms underlying conscious visual perception are often studied with either binocular rivalry or perceptual rivalry stimuli. Despite existing research into both types of rivalry, it remains unclear to what extent their underlying mechanisms involve common computational rules. Computational models of binocular rivalry mechanisms are generally tested against Levelt's four propositions, describing the psychophysical relation between stimulus strength and alternation dynamics in binocular rivalry. Here we use a bistable rotating structure-from-motion sphere, a generally studied form of perceptual rivalry, to demonstrate that Levelt's propositions also apply to the alternation dynamics of perceptual rivalry. Importantly, these findings suggest that bistability in structure-from-motion results from active cross-inhibition between neural populations with computational principles similar to those present in binocular rivalry. Thus, although the neural input to the computational mechanism of rivalry may stem from different cortical neurons and different cognitive levels the computational principles just prior to the production of visual awareness appear to be common to the two types of rivalry.

Visual rivalry is thought to be a distributed process that simultaneously takes place at multiple levels in the visual processing hierarchy. Also, the different types of rivalry, such as binocular and monocular rivalry, are thought to engage shared underlying mechanisms. We hypothesized that the amount of perceptual suppression during rivalry as measured by the total duration of fully exclusive perceptual dominance is determined by a distance in a neurally represented feature space. This hypothesis can be contrasted with the possibility that the brain constructs an internal model of the outside world using full-fledged object representations, and that perceptual suppression is due to an appraisal of the likelihood of the particular stimulus configuration at a high, object-based level. We applied color and stereo-depth differences between monocular rivalry stimulus gratings, and manipulated color and eye-of-origin information in binocular rivalry using the flicker & switch presentation paradigm. Our data show that exclusivity in visual rivalry increases with increased difference in feature space without regard for real-world constraints, and that eye-of-origin information may be regarded as a segregating feature that functions in a manner similar to color and stereo-depth information. Moreover, distances defined in multiple feature dimensions additively and independently increase the amount of perceptual exclusivity and coherence in both monocular and binocular rivalry. We conclude that exclusivity in visual rivalry is determined by a distance in feature space that is subtended by multiple stimulus features.

Two of the strongest tools to manipulate visual awareness of potentially salient stimuli are binocular rivalry and dichoptic masking. Binocular rivalry is induced by presenting incompatible images to the two eyes over prolonged periods of time, leading to an alternating perception of the two images. Dichoptic masking is induced when two images are presented once in rapid succession, leading to the perception of just one of the images. Although these phenomena share some key characteristics, most notably the ability to erase from awareness potentially very salient stimuli, their relationship is poorly understood. We investigated the perceptual dynamics during long-lasting dynamic stimulation leading to binocular rivalry or dichoptic masking. We show that the perceptual dynamics during dichoptic masking conditions meet the classifiers used to classify a process as binocular rivalry; that is, (1) Levelt's 2nd proposition is obeyed; (2) perceptual dominance durations follow a gamma distribution; and (3) dominance durations are sequentially independent. We suggest that binocular rivalry and dichoptic masking may be mediated by the same inhibitory mechanisms.

We show that previewing one half image of a binocular rivalry pair can cause it to gain initial dominance when the other half is added, a novel phenomenon we term flash facilitation. This is the converse of a known effect called flash suppression, where the previewed image becomes suppressed upon rivalrous presentation. The exact effect of previewing an image depends on both the duration and the contrast of the prior stimulus. Brief, low-contrast prior stimuli facilitate, whereas long, high-contrast ones suppress. These effects have both an eye-based component and a pattern-based component. Our results suggest that, instead of reflecting two unrelated mechanisms, both facilitation and suppression are manifestations of a single process that occurs progressively during presentation of the prior stimulus. The distinction between the two phenomena would then lie in the extent to which the process has developed during prior stimulation. This view is consistent with a neural model previously proposed to account for perceptual stabilization of ambiguous stimuli, suggesting a relation between perceptual stabilization and the present phenomena.

After viewing directional motion, one is likely to perceive a subsequently presented directionally ambiguous motion as being in the same direction as the prior motion. The perceptual bias towards the most recent percept gradually develops as the interval between the prior stimulus and a subsequent test becomes longer. This form of positive bias, or priming, is created in an automatic fashion. It remain unclear how such perceptual bias could be eliminated by a stimulus manipulation. Here we examine whether presentation of a stimulus, which was neutral as to the competing perceptual interpretations, during the interval between prior and test stimuli, disrupts the development of the priming effect. In experiments with ambiguous motion, we used stationary gratings as the neutral stimuli, and in an experiment with binocular rivalry between orthogonal gratings, we used a plaid pattern consisting of the two rival gratings. In both cases, presenting the neutral stimuli reduced the perceptual bias. These findings show that the visual system dynamically calibrates its internal bias using a recent percept and that this internal bias can be nullified by presenting neutral stimuli.

State transitions in the nervous system often take shape as traveling waves, whereby one neural state is replaced by another across space in a wave-like manner. In visual perception, transitions between the two mutually exclusive percepts that alternate when the two eyes view conflicting stimuli (binocular rivalry) may also take shape as traveling waves. The properties of these waves point to a neural substrate of binocular rivalry alternations that have the hallmark signs of lower cortical areas. In a series of experiments, we show a potent interaction between traveling waves in binocular rivalry and stimulus motion. The course of the traveling wave is biased in the motion direction of the suppressed stimulus that gains dominance by means of the wave-like transition. Thus, stimulus motion may propel the traveling wave across the stimulus to the extent that the stimulus motion dictates the traveling wave's direction completely. Using a computational model, we show that a speed-dependent asymmetry in lateral inhibitory connections between retinotopically organized and motion-sensitive neurons can explain our results. We argue that such a change in suppressive connections may play a vital role in the resolution of dynamic occlusion situations.

Recent work investigated the influence of exogenous attention on initial percept dominance at the onset of binocular rivalry. It was reported that cueing attention to one of two binocularly presented transparent stimuli immediately prior to rivalrous viewing provided the cued stimulus with a competitive advantage in subsequent binocular rivalry. This effect was independent of the eye containing the cued stimulus during the rivalry phase. In this recent work, the attention cue was always presented to both eyes. This leaves unclear the extent to which cueing affects binocular and/or monocular stimulus representations. To disambiguate this issue, we compared the cueing strength when the cue was presented ipsi-, contra- or bi-laterally with respect to the eye containing the cued stimulus during subsequent binocular rivalry. Besides replicating previous findings, we found that stimulus cueing readily transfers across eyes, suggesting that binocular mechanisms mediate exogenous attention effects on dominance selection at the onset of binocular rivalry.

We investigated the role of retinotopic visual cortex and motion-sensitive areas in representing the content of visual awareness during ambiguous structure-from-motion (SFM), using functional magnetic resonance imaging (fMRI) and multivariate statistics (support vector machines). Our results indicate that prediction of perceptual states can be very accurate for data taken from dorsal visual areas V3A, V4D, V7, and MT+ and for parietal areas responsive to SFM, but to a lesser extent for other visual areas. Generalization of prediction was possible, because prediction accuracy was significantly better than chance for both an unambiguous stimulus and a different experimental design. Detailed analysis of eye movements revealed that strategic and even encouraged beneficial eye movements were not the cause of the prediction accuracy based on cortical activation. We conclude that during perceptual rivalry, neural correlates of visual awareness can be found in retinotopic visual cortex, MT+, and parietal cortex. We argue that the organization of specific motion-sensitive neurons creates detectable biases in the preferred direction selectivity of voxels, allowing prediction of perceptual states. During perceptual rivalry, retinotopic visual cortex, in particular higher-tier dorsal areas like V3A and V7, actively represents the content the visual awareness.

When the two eyes are presented with sufficiently different stimuli, the stimuli will engage in binocular rivalry. During binocular rivalry, a subject's perceptual state alternates between awareness of the stimulus presented to the right eye and that presented to the left eye. There are instances in which competition is not eye-based, but instead takes place between stimulus features, as is the case in flicker and switch rivalry (F&S). Here we investigate another such instance, interocular grouping, using a Diaz-Caneja type stimulus in conjunction with synchronous stimulus flicker. Our results indicate that stimulus flicker increases the total duration of interocularly bound percepts, and that this effect occurs for a range of temporal flicker frequencies. Furthermore, the use of contrast-inversion flicker causes a decrease of total dominance duration of the interocularly bound percepts. We argue that different flickering regimes can be used to differentially stimulate lower and higher levels of visual processing involved in binocular rivalry. We propose that the amount of interocularly combined pattern-completed percept can be regarded as a measure of the level at which binocular rivalry is resolved.

Motion is fully described by a direction and a speed. The processing of direction information by the visual system has been extensively studied; much less is known, however, about the processing of speed. Although it is generally accepted that the direction of motion is processed by a single motion system, no such consensus exists for speed. Psychophysical data from humans suggest two separate systems processing luminance-based fast and slow speeds, whereas neurophysiological recordings in monkeys generally show continuous speed representation, hinting at a single system. Although the neurophysiological findings hint at a single system, they remain inconclusive as only a limited amount of cells can be measured per study and, possibly, the putative different motion systems are anatomically separate. In three psychophysical motion adaptation experiments, we show that predictions on the basis of the two-motion system hypothesis are not met. Instead, concurrent modeling showed that both here-presented and previous data are consistent with a single system subserving human speed perception. These findings have important implications for computational models of motion processing and the low-level organization of the process.

Any computation of metric surface structure from horizontal disparities depends on the viewing geometry, and analysing this dependence allows us to narrow down the choice of viable schemes. For example, all depth-based or slant-based schemes (i.e. nearly all existing models) are found to be unrealistically sensitive to natural errors in vergence. Curvature-based schemes avoid these problems and require only moderate, more robust view-dependent corrections to yield local object shape, without any depth coding. This fits the fact that humans are strikingly insensitive to global depth but accurate in discriminating surface curvature. The latter also excludes coding only affine structure. In view of new adaptation results, our goal becomes to directly extract retinotopic fields of metric surface curvatures (i.e. avoiding intermediate disparity curvature). To find a robust neural realisation, we combine new exact analysis with basic neural and psychophysical constraints. Systematic, step-by-step 'design' leads to neural operators which employ a novel family of 'dynamic' receptive fields (RFs), tuned to specific (bi-)local disparity structure. The required RF family is dictated by the non-Euclidean geometry that we identify as inherent in cyclopean vision. The dynamic RF-subfield patterns are controlled via gain modulation by binocular vergence and version, and parameterised by a cell-specific tuning to slant. Our full characterisation of the neural operators invites a range of new neurophysiological tests. Regarding shape perception, the model inverts widely accepted interpretations: It predicts the various types of errors that have often been mistaken for evidence against metric shape extraction.

We investigated the influence of changing physical parameters and task on bistable perception of an ambiguously rotating sphere (SFM). Increasing dot-density and velocity decreased the duration of perceptual phases during both passive viewing and voluntary control exertion. Our main finding is that voluntary control of perception depends on the physical parameters constituting the stimulus. This dependency places important constraints on the mechanisms mediating voluntary control as these mechanisms cannot operate independently of stimulus characteristics. In addition, local asymmetries in dot-densities can trigger alternations towards the most salient direction, which is not necessarily associated with largest number of dots: competition between perceptual interpretations during SFM appears to occur between surface-based representations rather than between individual elements. Finally, we show that voluntary control remains effective, even when attentive tracking of individual stimulus elements is no longer possible.

Although it is known that high-level spatial attention affects adaptation for a variety of stimulus features (including binocular disparity), the influence of voluntary attentional control-and the associated awareness-on adaptation has remained unexplored. We developed an ambiguous surface slant adaptation stimulus with conflicting monocular and binocular slant signals that instigated two mutually exclusive surface percepts with opposite slants. Using intermittent stimulus removal, subjects were able to voluntarily select one of the two rivaling slant percepts for extended adaptation periods, enabling us to dissociate slant adaptation due to awareness from stimulus-induced slant adaptation. We found that slant aftereffects (SAE) for monocular and binocular test patterns had opposite signs when measured simultaneously. There was no significant influence of voluntarily controlled perceptual state during adaptation on SAEs of monocular or binocular signals. In addition, the magnitude of the binocular SAE did not correlate with the magnitude of perceived slant. Using adaptation to one slant cue, and testing with the other cue, we demonstrated that multiple slant signals adapt independently. We conclude that slant adaptation occurs before the level of slant awareness. Our findings place the site of stereoscopic slant adaptation after disparity and eye posture are interpreted for slant [as demonstrated by Berends et al. (Berends, E. M., Liu, B., & Schor, C. M. (2005). Stereo-slant adaptation is high level and does not involve disparity coding. Journal of Vision 5 (1), 71-80), using that disparity scales with distance], but before other slant signals are integrated for the resulting awareness of the presented slant stimulus.

We studied distributions of perceptual rivalry reversals, as defined by the two fitted parameters of the Gamma distribution. We did so for a variety of bi-stable stimuli and voluntary control exertion tasks. Subjects' distributions differed from one another for a particular stimulus and control task in a systematic way that reflects a constraint on the describing parameters. We found a variety of two-parameter effects, the most important one being that distributions of subjects differ from one another in the same systematic way across different stimuli and control tasks (i.e., a fast switcher remains fast across all conditions in a parameter-specified way). The cardinal component of subject-dependent variation was not the conventionally used mean reversal rate, but a component that was oriented-for all stimuli and tasks-roughly perpendicular to the mean rate. For the Necker cube, we performed additional experiments employing specific variations in control exertion, suggesting that subjects have to a considerable extent independent control over the reversal rate of either of the two competing percepts.

We have investigated the role of saccades and fixation positions in two perceptual rivalry paradigms (slant rivalry and Necker cube) and in two binocular rivalry paradigms (grating and house-face rivalry), and we compared results obtained from two different voluntary control conditions (natural viewing and hold percept). We found that for binocular rivalry, rather than for perceptual rivalry, there is a marked positive temporal correlation between saccades and perceptual flips at about the moment of the flip. Across different voluntary control conditions the pattern of temporal correlation did not change (although the amount of correlation did frequently, but not always, change), indicating that subjects do not use different temporal eye movement schemes to exert voluntary control. Analysis of the fixation positions at about the moment of the flips indicates that the fixation position by itself does not determine the percept but that subjects prefer to fixate at different positions when asked to hold either of the different percepts.

Using event-related functional magnetic resonance imaging, we studied the activation correlating with the awareness of stereoscopic depth using a bistable slanted surface (slant rivalry). Bistability resulted from incongruence between two slant-defining cues: binocular disparity and monocular perspective. The stimulus was perceived as alternating between the perspective-dominated percept (monocular depth) and the disparity-dominated percept (stereopsis), while sensory input remained constant, enabling us to study changes in awareness of depth associated with either cue. Transient activation relating to perceptual alternations was found bilaterally in the caudal part of the intraparietal sulcus, in the right-hemispheric anterior intraparietal sulcus, within visual area V4d-topo, and inferior to area MT+. Transient activation correlating specifically with alternations toward the disparity-dominated percept was found in a number of visual areas, including dorsal visual areas V3A, V7, and V4d-topo and visual areas MT+ and lateral occipital complex. No activation was found for alternations toward the perspective-dominated percept. Our results show that of all visual areas responsive to disparity-defined depth, V4d-topo shows the most robust signal changes correlating with the instigation of stereoscopic depth awareness (stereopsis).