Perceptual Switching Research Articles

Perceptual switches during structure-from-motion do not elicit pupil response

Perceptual switches during structure-from-motion do not elicit pupil response

Spontaneous Eye Blinks Map the Probability of Perceptual Reinterpretation During Visual and Auditory Ambiguity.

Spontaneous eye blinks are modulated around perceptual events. Our previous study, using a visual ambiguous stimulus, indicated that blink probability decreases before a reported perceptual switch. In the current study, we tested our hypothesis that an absence of blinks marks a time in which perceptual switches are facilitated in- and outside the visual domain. In three experiments, presenting either a visual motion quartet in light or darkness or a bistable auditory streaming stimulus, we found a co-occurrence of blink rate reduction with increased perceptual switch probability. In the visual domain, perceptual switches induced by a short interruption of visual input (blank) allowed an estimate of the timing of the perceptual event with respect to the motor response. This provided the first evidence that the blink reduction was not a consequence of the perceptual switch. Importantly, by showing that the time between switches and the previous blink was significantly longer than the inter-blink interval, our studies allowed to conclude that perceptual switches did not happen at random but followed a prolonged period of nonblinking. Correspondingly, blink rate and switch rate showed an inverse relationship. Our study supports the idea that the absence or presence of blinks maps perceptual processes independent of the sensory modality.

An Accumulating Neural Signal Underlying Binocular Rivalry Dynamics.

During binocular rivalry, conflicting images are presented one to each eye and perception alternates stochastically between them. Despite stable percepts between alternations, modeling suggests that neural signals representing the two images change gradually, and that the duration of stable percepts are determined by the time required for these signals to reach a threshold that triggers an alternation. However, direct physiological evidence for such signals has been lacking. Here, we identify a neural signal in the human visual cortex that shows these predicted properties. We measured steady-state visual evoked potentials (SSVEPs) in 84 human participants (62 females, 22 males) who were presented with orthogonal gratings, one to each eye, flickering at different frequencies. Participants indicated their percept while EEG data were collected. The time courses of the SSVEP amplitudes at the two frequencies were then compared across different percept durations, within participants. For all durations, the amplitude of signals corresponding to the suppressed stimulus increased and the amplitude corresponding to the dominant stimulus decreased throughout the percept. Critically, longer percepts were characterized by more gradual increases in the suppressed signal and more gradual decreases of the dominant signal. Changes in signals were similar and rapid at the end of all percepts, presumably reflecting perceptual transitions. These features of the SSVEP time courses are well predicted by a model in which perceptual transitions are produced by the accumulation of noisy signals. Identification of this signal underlying binocular rivalry should allow strong tests of neural models of rivalry, bistable perception, and neural suppression.SIGNIFICANCE STATEMENT During binocular rivalry, two conflicting images are presented to the two eyes and perception alternates between them, with switches occurring at seemingly random times. Rivalry is an important and longstanding model system in neuroscience, used for understanding neural suppression, intrinsic neural dynamics, and even the neural correlates of consciousness. All models of rivalry propose that it depends on gradually changing neural activity that on reaching some threshold triggers the perceptual switches. This manuscript reports the first physiological measurement of neural signals with that set of properties in human participants. The signals, measured with EEG in human observers, closely match the predictions of recent models of rivalry, and should pave the way for much future work.

Studying perceptual bias in favor of the from-above Necker cube perspective in a goal-directed behavior.

When viewing a completely ambiguous image, different interpretations can switch involuntarily due to internal top-down processing. In the case of the Necker cube, an entirely ambiguous stimulus, observers often display a bias in perceptual switching between two interpretations based on their perspectives: one with a from-above perspective (FA) and the other with a from-below perspective (FB). Typically, observers exhibit a priori top-down bias in favor of the FA interpretation, which may stem from a statistical tendency in everyday life where we more frequently observe objects from above. However, it remains unclear whether this perceptual bias persists when individuals voluntarily decide on the Necker cube's interpretation in goal-directed behavior, and the impact of ambiguity in this context is not well-understood. In our study, we instructed observers to voluntarily identify the orientation of a Necker cube while manipulating its ambiguity from low (LA) to high (HA). Our investigation aimed to test two hypotheses: (i) whether the perspective (FA or FB) would result in a bias in response time, and (ii) whether this bias would depend on the level of stimulus ambiguity. Additionally, we analyzed electroencephalogram (EEG) signals to identify potential biomarkers that could explain the observed perceptual bias. The behavioral results confirmed a perceptual bias in favor of the from-above perspective, as indicated by shorter response times. However, this bias diminished for stimuli with high ambiguity. For the LA stimuli, the occipital theta-band power consistently exceeded the frontal theta-band power throughout most of the decision-making time. In contrast, for the HA stimuli, the frontal theta-band power started to exceed the occipital theta-band power during the 0.3-s period preceding the decision. We propose that occipital theta-band power reflects evidence accumulation, while frontal theta-band power reflects its evaluation and decision-making processes. For the FB perspective, occipital theta-band power exhibited higher values and dominated over a longer duration, leading to an overall increase in response time. These results suggest that more information and more time are needed to encode stimuli with a FB perspective, as this template is less common for the observers compared to the template for a cube with a FA perspective.

Neural correlates of perceptual switching and their association with empathy and alexithymia in individuals with and without autism spectrum disorder

Individuals with autism spectrum disorder (ASD) often show limited empathy (poor recognition of others’ emotions) and high alexithymia (poor recognition of own emotions and external thinking), which can negatively impact their social functioning. Previous experimental studies suggest that alterations in cognitive flexibility play key roles in the development of these characteristics in ASD. However, the underlying neural mechanisms that link cognitive flexibility and empathy/alexithymia are still largely unknown. In this study, we examined the neural correlates of cognitive flexibility via functional magnetic resonance imaging during perceptual task-switching in typical development (TD) adults and adults with ASD. We also investigated associations between regional neural activity and psychometric empathy and alexithymia scores among these populations. In the TD group, stronger activation of the left middle frontal gyrus was associated with better perceptual switching and greater empathic concern. Among individuals with ASD, stronger activation of the left inferior frontal gyrus was associated with better perceptual switching, greater empathy, and lower alexithymia. These findings will contribute to develop a better understanding of social cognition, and could be informative for the development of new ASD therapies.

Attractor-Like Dynamics Extracted from Human Electrocorticographic Recordings Underlie Computational Principles of Auditory Bistable Perception.

In bistable perception, observers experience alternations between two interpretations of an unchanging stimulus. Neurophysiological studies of bistable perception typically partition neural measurements into stimulus-based epochs and assess neuronal differences between epochs based on subjects' perceptual reports. Computational studies replicate statistical properties of percept durations with modeling principles like competitive attractors or Bayesian inference. However, bridging neuro-behavioral findings with modeling theory requires the analysis of single-trial dynamic data. Here, we propose an algorithm for extracting nonstationary timeseries features from single-trial electrocorticography (ECoG) data. We applied the proposed algorithm to 5-min ECoG recordings from human primary auditory cortex obtained during perceptual alternations in an auditory triplet streaming task (six subjects: four male, two female). We report two ensembles of emergent neuronal features in all trial blocks. One ensemble consists of periodic functions that encode a stereotypical response to the stimulus. The other comprises more transient features and encodes dynamics associated with bistable perception at multiple time scales: minutes (within-trial alternations), seconds (duration of individual percepts), and milliseconds (switches between percepts). Within the second ensemble, we identified a slowly drifting rhythm that correlates with the perceptual states and several oscillators with phase shifts near perceptual switches. Projections of single-trial ECoG data onto these features establish low-dimensional attractor-like geometric structures invariant across subjects and stimulus types. These findings provide supporting neural evidence for computational models with oscillatory-driven attractor-based principles. The feature extraction techniques described here generalize across recording modality and are appropriate when hypothesized low-dimensional dynamics characterize an underlying neural system.SIGNIFICANCE STATEMENT Irrespective of the sensory modality, neurophysiological studies of multistable perception have typically investigated events time-locked to the perceptual switching rather than the time course of the perceptual states per se. Here, we propose an algorithm that extracts neuronal features of bistable auditory perception from largescale single-trial data while remaining agnostic to the subject's perceptual reports. The algorithm captures the dynamics of perception at multiple timescales, minutes (within-trial alternations), seconds (durations of individual percepts), and milliseconds (timing of switches), and distinguishes attributes of neural encoding of the stimulus from those encoding the perceptual states. Finally, our analysis identifies a set of latent variables that exhibit alternating dynamics along a low-dimensional manifold, similar to trajectories in attractor-based models for perceptual bistability.

Altered visual conscious awareness in patients with vestibular dysfunctions; a cross-sectional observation study

BackgroundPatients with vestibular dysfunctions often experience visual-induced symptoms. Here we asked whether such visual dependence can be related to alterations in visual conscious awareness in these patients. MethodsTo measure visual conscious awareness, we used the effect of motion-induced blindness (MIB,) in which the perceptual awareness of the visual stimulus alternates despite its unchanged physical characteristics. In this phenomenon, a salient visual target spontaneously disappears and subsequently reappears from visual perception when presented against a moving visual background. The number of perceptual switches during the experience of the MIB stimulus was measured for 120 s in 15 healthy controls, 15 patients with vestibular migraine, 15 patients with benign positional paroxysmal vertigo (BPPV) and 15 with migraine without vestibular symptoms. ResultsPatients with vestibular dysfunctions (i.e., both vestibular migraine and BPPV) exhibited increased perceptual fluctuations during MIB compared to healthy controls and migraine patients without vertigo. In VM patients, those with more severe symptoms exhibited higher fluctuations of visual awareness (i.e., positive correlation), whereas, in BPPV patients, those with more severe symptoms had lower fluctuations of visual awareness (i.e., negative correlation). ImplicationsTaken together, these findings show that fluctuations of visual awareness are linked to the severity of visual-induced symptoms in patients with vestibular dysfunctions, and distinct pathophysiological mechanisms may mediate visual vertigo in peripheral versus central vestibular dysfunctions.

Positive hysteresis in emotion recognition: Face processing visual regions are involved in perceptual persistence, which mediates interactions between anterior insula and medial prefrontal cortex

Facial emotion perception can be studied from the point of view of dynamic systems whose output may depend not only on current input but also on prior history — a phenomenon known as hysteresis. In cognitive neuroscience, hysteresis has been described as positive (perceptual persistence) or negative (fatigue of current percept) depending on whether perceptual switching occurs later or earlier than actual physical stimulus changes. However, its neural correlates remain elusive. We used dynamic transitions between emotional expressions and combined behavioral assessment with functional magnetic resonance imaging (fMRI) to investigate the underlying circuitry of perceptual hysteresis in facial emotion recognition. Our findings revealed the involvement of face-selective visual areas — fusiform face area (FFA) and superior temporal sulcus (STS) — in perceptual persistence as well as the right anterior insula. Moreover, functional connectivity analyses revealed an interplay between the right anterior insula and medial prefrontal cortex, which showed to be dependent on the presence of positive hysteresis. Our results support the hypothesis that high-order regions are involved in perceptual stabilization and decision during perceptual persistence (positive hysteresis) and add evidence to the role of the anterior insula as a hub of sensory information in perceptual decision-making.

A wrinkle in and of time: Contraction of felt duration with a single perceptual switch

The way we represent and perceive time has crucial implications for studying temporality in conscious experience. Contrasting positions posit that temporal information is separately abstracted out like any other perceptual property through specialized mechanisms or that time is represented through the temporality of experiences themselves. To add to this debate, we investigate alterations in felt time in conditions where only conscious visual experience is altered through perceptual switches while a bistable figure remains physically unchanged. We predicted that if perceived time is a function of temporally evolving conscious content, then a break in it (here via a perceptual switch) would also lead to a break in felt time. In three experiments, we showed participants a Necker cube that was manipulated to induce a perceptual switch (experiments 1(a) and 1(b)) or left to switch on its own (experiment 2). We asked participants to report both perceptual switches and felt durations (experiments 1(a) and 2) or only estimate time (experiment 1(b)). Over these three experiments, we find evidence of contraction of felt time in trials with a perceptual switch, consistent with the idea that perceived time is a function of temporally evolving conscious experience. Additionally, we present a phenomenological demonstration to support our empirical data. Overall, the study provides evidence for temporal mirroring and isomorphism in visual experience, arguing for a link between the timing of experience and time perception.

Neural oscillations promoting perceptual stability and perceptual memory during bistable perception

Ambiguous images elicit bistable perception, wherein periods of momentary perceptual stability are interrupted by sudden perceptual switches. When intermittently presented, ambiguous images trigger a perceptual memory trace in the intervening blank periods. Understanding the neural bases of perceptual stability and perceptual memory during bistable perception may hold clues for explaining the apparent stability of visual experience in the natural world, where ambiguous and fleeting images are prevalent. Motivated by recent work showing the involvement of the right inferior frontal gyrus (rIFG) in bistable perception, we conducted a transcranial direct-current stimulation (tDCS) study with a double-blind, within-subject cross-over design to test a potential causal role of rIFG in these processes. Subjects viewed ambiguous images presented continuously or intermittently while under EEG recording. We did not find any significant tDCS effect on perceptual behavior. However, the fluctuations of oscillatory power in the alpha and beta bands predicted perceptual stability, with higher power corresponding to longer percept durations. In addition, higher alpha and beta power predicted enhanced perceptual memory during intermittent viewing. These results reveal a unified neurophysiological mechanism sustaining perceptual stability and perceptual memory when the visual system is faced with ambiguous input.

Conflict monitoring and attentional adjustment during binocular rivalry.

To make sense of ambiguous and, at times, fragmentary sensory input, the brain must rely on a process of active interpretation. At any given moment, only one of several possible perceptual representations prevails in our conscious experience. Our hypothesis is that the competition between alternative representations induces a pattern of neural activation resembling cognitive conflict, eventually leading to fluctuations between different perceptual outcomes in the case of steep competition. To test this hypothesis, we probed changes in perceptual awareness between competing images using binocular rivalry. We drew our predictions from the conflict monitoring theory, which holds that cognitive control is invoked by the detection of conflict during information processing. Our results show that fronto-medial theta oscillations (5-7 Hz), an established electroencephalography (EEG) marker of conflict, increases right before perceptual alternations and decreases thereafter, suggesting that conflict monitoring occurs during perceptual competition. Furthermore, to investigate conflict resolution via attentional engagement, we looked for a neural marker of perceptual switches as by parieto-occipital alpha oscillations (8-12 Hz). The power of parieto-occipital alpha displayed an inverse pattern to that of fronto-medial theta, reflecting periods of high interocular inhibition during stable perception, and low inhibition around moments of perceptual change. Our findings aim to elucidate the relationship between conflict monitoring mechanisms and perceptual awareness.

Long-term priors influence visual perception through recruitment of long-range feedback

Perception results from the interplay of sensory input and prior knowledge. Despite behavioral evidence that long-term priors powerfully shape perception, the neural mechanisms underlying these interactions remain poorly understood. We obtained direct cortical recordings in neurosurgical patients as they viewed ambiguous images that elicit constant perceptual switching. We observe top-down influences from the temporal to occipital cortex, during the preferred percept that is congruent with the long-term prior. By contrast, stronger feedforward drive is observed during the non-preferred percept, consistent with a prediction error signal. A computational model based on hierarchical predictive coding and attractor networks reproduces all key experimental findings. These results suggest a pattern of large-scale information flow change underlying long-term priors’ influence on perception and provide constraints on theories about long-term priors’ influence on perception.

Causal roles of prefrontal cortex during spontaneous perceptual switching are determined by brain state dynamics.

The prefrontal cortex (PFC) is thought to orchestrate cognitive dynamics. However, in tests of bistable visual perception, no direct evidence supporting such presumable causal roles of the PFC has been reported except for a recent work. Here, using a novel brain-state-dependent neural stimulation system, we identified causal effects on percept dynamics in three PFC activities-right frontal eye fields, dorsolateral PFC (DLPFC), and inferior frontal cortex (IFC). The causality is behaviourally detectable only when we track brain state dynamics and modulate the PFC activity in brain-state-/state-history-dependent manners. The behavioural effects are underpinned by transient neural changes in the brain state dynamics, and such neural effects are quantitatively explainable by structural transformations of the hypothetical energy landscapes. Moreover, these findings indicate distinct functions of the three PFC areas: in particular, the DLPFC enhances the integration of two PFC-active brain states, whereas IFC promotes the functional segregation between them. This work resolves the controversy over the PFC roles in spontaneous perceptual switching and underlines brain state dynamics in fine investigations of brain-behaviour causality.

Pupil constriction via the parasympathetic pathway precedes perceptual switch of ambiguous stimuli

Perceptual rivalry of ambiguous stimuli reflects the interaction of neural activity among multiple cortical regions. However, it remains unclear what drives a spontaneous perceptual alteration. We hypothesized that increased fluctuations in spontaneous neural activity due to arousal reduction drive the perceptual switch. Here, we show that the pupils shrank a few seconds prior to the onset of the spontaneous perceptual switch. Such pupil constriction was not observed before the exogenous perceptual switch. Pharmacological experiments confirmed that the pupil constriction disappeared when the peripheral parasympathetic pathway (pupil sphincter muscle) was blocked, but it remained intact when the peripheral sympathetic pathway (pupil dilator muscle) was manipulated. Furthermore, rapid pupil dilations with behavioral response are also mediated by the peripheral parasympathetic pathway. The present findings suggested that transient arousal drops, as denoted by the autonomic nervous modulation of pupil size, are involved in inducing the spontaneous perceptual switch of bistable stimuli.

Pupillometry in auditory multistability.

In multistability, a constant stimulus induces alternating perceptual interpretations. For many forms of visual multistability, the transition from one interpretation to another (“perceptual switch”) is accompanied by a dilation of the pupil. Here we ask whether the same holds for auditory multistability, specifically auditory streaming. Two tones were played in alternation, yielding four distinct interpretations: the tones can be perceived as one integrated percept (single sound source), or as segregated with either tone or both tones in the foreground. We found that the pupil dilates significantly around the time a perceptual switch is reported (“multistable condition”). When participants instead responded to actual stimulus changes that closely mimicked the multistable perceptual experience (“replay condition”), the pupil dilated more around such responses than in multistability. This still held when data were corrected for the pupil response to the stimulus change as such. Hence, active responses to an exogeneous stimulus change trigger a stronger or temporally more confined pupil dilation than responses to an endogenous perceptual switch. In another condition, participants randomly pressed the buttons used for reporting multistability. In Study 1, this “random condition” failed to sufficiently mimic the temporal pattern of multistability. By adapting the instructions, in Study 2 we obtained a response pattern more similar to the multistable condition. In this case, the pupil dilated significantly around the random button presses. Albeit numerically smaller, this pupil response was not significantly different from the multistable condition. While there are several possible explanations–related, e.g., to the decision to respond–this underlines the difficulty to isolate a purely perceptual effect in multistability. Our data extend previous findings from visual to auditory multistability. They highlight methodological challenges in interpreting such data and suggest possible approaches to meet them, including a novel stimulus to simulate the experience of perceptual switches in auditory streaming.

The Role of Blinks, Microsaccades and their Retinal Consequences in Bistable Motion Perception.

Eye-related movements such as blinks and microsaccades are modulated during bistable perceptual tasks. However, if they play an active role during internal perceptual switches is not known. We conducted two experiments involving an ambiguous plaid stimulus, wherein participants were asked to continuously report their percept, which could consist of either unidirectional coherent or bidirectional component movement. Our main results show that blinks and microsaccades did not facilitate perceptual switches. On the contrary, a reduction in eye movements preceded the perceptual switch. Blanks, on the other hand, thought to mimic the retinal consequences of a blink, consistently led to a switch. Through the timing of the blank-introduced perceptual change, we were able to estimate the delay between the internal switch and the response. This delay further allowed us to evaluate that the reduction in blink probability co-occurred with the internal perceptual switch. Additionally, our results indicate that distinct internal processes underlie the switch to coherent vs. component percept. Blanks exclusively facilitated a switch to the coherent percept, and only the switch to coherent percept was followed by an increase in blink rate. In a second study, we largely replicated the findings and included a microsaccade analysis. Microsaccades only showed a weak relation with perceptual switches, but their direction was correlated with the perceived motion direction. Nevertheless, our data suggests an interaction between microsaccades and blinks by showing that microsaccades were differently modulated around blinks compared with blanks. This study shows that a reduction in eye movements precedes internal perceptual switches indicating that the rate of blinks can set the stage for a reinterpretation of sensory input. While a perceptual switch based on changed sensory input usually leads to an increase in blink rate, such an increase was only present after the perceptual switch to coherent motion but absent after the switch to component percept. This provides evidence of different underlying mechanism or internal consequence of the two perceptual switches and suggests that blinks can uncover differences in internal percept-related processes that are not evident from the percept itself.

Divergent Thinking Influences the Perception of Ambiguous Visual Illusions.

This study investigated the relationships between personality and creativity in the perception of two different ambiguous visual illusions. Previous research has suggested that Industriousness and Openness/Intellect (as measured by the Big Five Aspects Scale) are both associated with individual differences in perceptual switching rates for binocular rivalry stimuli. Here, we examined whether these relationships generalise to the Necker Cube and the Spinning Dancer illusions. In the experimental phase of this study, participants viewed these ambiguous figures under both static and dynamic, as well as free-view and fixation, conditions. As predicted, perceptual switching rates were higher: (a) for the static Necker Cube than the Spinning Dancer, and (b) in free-view compared with fixation conditions. In the second phase of the study, personality type and divergent thinking were measured using the Big Five Aspects Scale and the Alternate Uses Task, respectively. Higher creativity/divergent thinking (as measured by the Alternate Uses Task) was found to predict greater switching rates for the static Necker Cube (but not the Spinning Dancer) under both free-view and fixation conditions. These findings suggest that there are differences in the perceptual processing of creative individuals.

Individual Differences in Perceptual Organization: Reanalyzing Thurstone’s classic (1944) data and rediscovering factors for geometrical illusions, perceptual switching, and holistic ‘Gestalt’ closure

Individual Differences in Perceptual Organization: Reanalyzing Thurstone’s classic (1944) data and rediscovering factors for geometrical illusions, perceptual switching, and holistic ‘Gestalt’ closure

Perceptual switch creates a transient bias in favor of the new state at neighboring locations. Stimulus ambiguity does not matter.

Perceptual switch creates a transient bias in favor of the new state at neighboring locations. Stimulus ambiguity does not matter.

Meditators Exercise Better Endogenous and Exogenous Control of Visual Awareness

Meditation practice has shown improvements in perception, attention, and cognitive control. Here, we study the extent of endogenous and exogenous control meditators have over their visual awareness, when they view a bi-stable image. In experiment one, we investigate the differences between meditators and controls in terms of endogenous perceptual control, i.e., when they are instructed to speed up or slow down perceptual switches when viewing a Necker cube. In experiment two, we investigate how stimulus context and flankers (exogenous perceptual control) affect dwell times of visual percepts for meditators and non-meditators. Our results show that meditators (in comparison with controls) have longer dwell times in the neutral baseline condition. Meditators are also able to speed up and slow down the dwell times of their percepts more (relative to their baseline) than non-meditators. The dwell time distributions of meditators (mean, variance, and skewness) are also immune to flanker effects while viewing bi-stable images. The results show that meditators have better endogenous (instruction driven) and exogenous (stimulus driven) control compared with non-meditators over the contents and properties of their perceptual experience. The control exhibited by meditators is critical for not only understanding the effects of meditation but points to the need for models of visual perception and cognitive control to incorporate those effects.