Depth perception from pairs of overlapping cues in pictorial displays

The experiments reported herein probe the visual cortical mechanisms that control near-far percepts in response to two-dimensional stimuli. Figural contrast is found to be a principal factor for the emergence of percepts of near versus far in pictorial stimuli, especially when stimulus duration is brief. Pictorial factors such as interposition (Experiment 1) and partial occlusion (Experiments 2 and 3) may cooperate, as generally predicted by cue combination models, or compete with contrast factors in the manner predicted by the FACADE model. In particular, if the geometrical configuration of an image favors activation of cortical bipole grouping cells, as at the top of a T-junction, then this advantage can cooperate with the contrast of the configuration to facilitate a near-far percept at a lower contrast than at an X-junction. Varying the exposure duration of the stimuli shows that the more balanced bipole competition in the X-junction case takes longer exposures to resolve than the bipole competition in the T-junction case (Experiment 3).     

Predictive properties of visual adaptation

What humans perceive depends in part on what they have previously experienced. After repeated exposure to one stimulus, adaptation takes place in the form of a negative correlation between the current percept and the last displayed stimuli. Previous work has shown that this negative dependence can extend to a few minutes in the past, but the precise extent and nature of the dependence in vision is still unknown. In two experiments based on orientation judgments, we reveal a positive dependence of a visual percept with stimuli presented remotely in the past, unexpectedly and in contrast to what is known for the recent past. Previous theories of adaptation have postulated that the visual system attempts to calibrate itself relative to an ideal norm or to the recent past. We propose instead that the remote past is used to estimate the world's statistics and that this estimate becomes the reference. According to this new framework, adaptation is predictive: the most likely forthcoming percept is the one that helps the statistics of the most recent percepts match that of the remote past.     

Duality in Binocular Rivalry: Distinct Sensitivity of Percept Sequence and Percept Duration to Imbalance between Monocular Stimuli

Background

Visual perception is usually stable and accurate. However, when the two eyes are simultaneously presented with conflicting stimuli, perception falls into a sequence of spontaneous alternations, switching between one stimulus and the other every few seconds. Known as binocular rivalry, this visual illusion decouples subjective experience from physical stimulation and provides a unique opportunity to study the neural correlates of consciousness. The temporal properties of this alternating perception have been intensively investigated for decades, yet the relationship between two fundamental properties - the sequence of percepts and the duration of each percept - remains largely unexplored.

Methodology/Principal Findings

Here we examine the relationship between the percept sequence and the percept duration by quantifying their sensitivity to the strength imbalance between two monocular stimuli. We found that the percept sequence is far more susceptible to the stimulus imbalance than does the percept duration. The percept sequence always begins with the stronger stimulus, even when the stimulus imbalance is too weak to cause a significant bias in the percept duration. Therefore, introducing a small stimulus imbalance affects the percept sequence, whereas increasing the imbalance affects the percept duration, but not vice versa. To investigate why the percept sequence is so vulnerable to the stimulus imbalance, we further measured the interval between the stimulus onset and the first percept, during which subjects experienced the fusion of two monocular stimuli. We found that this interval is dramatically shortened with increased stimulus imbalance.

Conclusions/Significance

Our study shows that in binocular rivalry, the strength imblanace between monocular stimuli has a much greater impact on the percept sequence than on the percept duration, and increasing this imbalance can accelerate the process responsible for the percept sequence.     

Multi-timescale perceptual history resolves visual ambiguity

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.     

BOLD Signal in Both Ipsilateral and Contralateral Retinotopic Cortex Modulates with Perceptual Fading

Under conditions of visual fixation, perceptual fading occurs when a stationary object, though present in the world and continually casting light upon the retina, vanishes from visual consciousness. The neural correlates of the consciousness of such an object will presumably modulate in activity with the onset and cessation of perceptual fading.

Method

In order to localize the neural correlates of perceptual fading, a green disk that had been individually set to be equiluminant with the orange background, was presented in one of the four visual quadrants; Subjects indicated with a button press whether or not the disk was subjectively visible as it perceptually faded in and out.

Results

Blood oxygen-level dependent (BOLD) signal in V1 and ventral retinotopic areas V2v and V3v decreases when the disk subjectively disappears, and increases when it subjectively reappears. This effect occurs in early visual areas both ipsilaterally and contralaterally to the fading figure. That is, it occurs regardless of whether the fading stimulus is presented inside or outside of the corresponding portion of visual field. In addition, we find that the microsaccade rate rises before and after perceptual transitions from not seeing to seeing the disk, and decreases before perceptual transitions from seeing to not seeing the disk. These BOLD signal changes could be driven by a global process that operates across contralateral and ipsilateral visual cortex or by a confounding factor, such as microsaccade rate.     

The Spatial Properties of L- and M-Cone Inputs to Electroretinograms That Reflect Different Types of Post-Receptoral Processing

We studied the spatial arrangement of L- and M-cone driven electroretinograms (ERGs) reflecting the activity of magno- and parvocellular pathways. L- and M-cone isolating sine wave stimuli were created with a four primary LED stimulator using triple silent substitution paradigms. Temporal frequencies were 8 and 12 Hz, to reflect cone opponent activity, and 30, 36 and 48 Hz to reflect luminance activity. The responses were measured for full-field stimuli and for different circular and annular stimuli. The ERG data confirm the presence of two different mechanisms at intermediate and high temporal frequencies. The responses measured at high temporal frequencies strongly depended upon spatial stimulus configuration. In the full-field conditions, the L-cone driven responses were substantially larger than the full-field M-cone driven responses and also than the L-cone driven responses with smaller stimuli. The M-cone driven responses at full-field and with 70° diameter stimuli displayed similar amplitudes. The L- and M-cone driven responses measured at 8 and 12 Hz were of similar amplitude and approximately in counter-phase. The amplitudes were constant for most stimulus configurations. The results indicate that, when the ERG reflects luminance activity, it is positively correlated with stimulus size. Beyond 35° retinal eccentricity, the retina mainly contains L-cones. Small stimuli are sufficient to obtain maximal ERGs at low temporal frequencies where the ERGs are also sensitive to cone-opponent processing.     

Audiovisual integration: an investigation of the "streaming-bouncing" phenomenon

Temporal aspects of the perceptual integration of audiovisual information were investigated by utilizing the visual "streaming-bouncing" phenomenon. When two identical visual objects move towards each other, coincide, and then move away from each other, the objects can either be seen as streaming past one another or bouncing off each other. Although the streaming percept is dominant, the bouncing percept can be induced by presenting an auditory stimulus during the visual coincidence of the moving objects. Here we show that the bounce-inducing effect of the auditory stimulus is paramount when its onset and offset occur in temporal proximity of the onset and offset of the period of visual coincidence of the moving objects. When the duration of the auditory stimulus exceeded this period, visual bouncing disappears. Implications for a temporal window of audiovisual integration and the design of effective audiovisual warning signals are discussed.     

Non-linear population firing rates and voltage sensitive dye signals in visual areas 17 and 18 to short duration stimuli

Visual stimuli of short duration seem to persist longer after the stimulus offset than stimuli of longer duration. This visual persistence must have a physiological explanation. In ferrets exposed to stimuli of different durations we measured the relative changes in the membrane potentials with a voltage sensitive dye and the action potentials of populations of neurons in the upper layers of areas 17 and 18. For durations less than 100 ms, the timing and amplitude of the firing and membrane potentials showed several non-linear effects. The ON response became truncated, the OFF response progressively reduced, and the timing of the OFF responses progressively delayed the shorter the stimulus duration. The offset of the stimulus elicited a sudden and strong negativity in the time derivative of the dye signal. All these non-linearities could be explained by the stimulus offset inducing a sudden inhibition in layers II-III as indicated by the strongly negative time derivative of the dye signal. Despite the non-linear behavior of the layer II-III neurons the sum of the action potentials, integrated from the peak of the ON response to the peak of the OFF response, was almost linearly related to the stimulus duration.     

A comprehensive model of audiovisual perception: both percept and temporal dynamics

The sparse information captured by the sensory systems is used by the brain to apprehend the environment, for example, to spatially locate the source of audiovisual stimuli. This is an ill-posed inverse problem whose inherent uncertainty can be solved by jointly processing the information, as well as introducing constraints during this process, on the way this multisensory information is handled. This process and its result--the percept--depend on the contextual conditions perception takes place in. To date, perception has been investigated and modeled on the basis of either one of two of its dimensions: the percept or the temporal dynamics of the process. Here, we extend our previously proposed audiovisual perception model to predict both these dimensions to capture the phenomenon as a whole. Starting from a behavioral analysis, we use a data-driven approach to elicit a bayesian network which infers the different percepts and dynamics of the process. Context-specific independence analyses enable us to use the model's structure to directly explore how different contexts affect the way subjects handle the same available information. Hence, we establish that, while the percepts yielded by a unisensory stimulus or by the non-fusion of multisensory stimuli may be similar, they result from different processes, as shown by their differing temporal dynamics. Moreover, our model predicts the impact of bottom-up (stimulus driven) factors as well as of top-down factors (induced by instruction manipulation) on both the perception process and the percept itself.     

Brightness versus apparent contrast. 1: Incremental and decremental disks with varying diameter.

This study describes the matching of the brightness and of the apparent contrast of foveal disks, presented as an increment or decrement with varying diameter against a 300 cd.m-2 background. If the brightness in the centre of the disks is matched with a constant reference brightness, the well-known spatial Broca-Sulzer phenomenon is obtained. This effect is not found if the apparent contrast of the disks is matched instead. All matching results and detection thresholds indicate that luminance increments and decrements are processed asymmetrically by the visual system: for decrements the threshold curve and curves of isobrightness as well as apparent iso-contrast are all shifted towards larger disk radii relative to those for increments. Two nonlinear single-channel models are tested against the data.     

The persistences of vision

Human observers continue to experience a visual stimulus for some time after the offset that stimulus. The neural activity evoked by a visual stimulus continues for some time after its offset. The information extracted from a visual stimulus continues to be registered in a visual form of memory ('iconic memory') for some time after its offset. We may thus distinguish three distinct senses in which a visual stimulus may be said to persist after its physical offset: there is phenomenological persistence, neural persistence and informational persistence. Various assumptions have been made about the relation between these forms of visual persistence. The most frequent assumption is that they correspond simply to three different methods for studying a single entity. Detailed consideration of what is known about the properties of these three forms of persistence suggests, however, that this assumption is not correct. It can reasonably be proposed that visible persistence is the phenomenological correlate of neural persistence occurring at various stages of the visual system: photoreceptors, ganglion cells and the stereopsis system. Iconic memory on the other hand, does not correspond to visible persistence, nor to neural persistence in any stage of the visual system. Recent work, in fact, suggests that iconic memory is a property of some relatively late stage in the visual information-processing system, rather than being a peripheral sensory buffer store. This suggestion raises some fundamental theoretical issues concerning the psychology of visual perception, issues with which cognitive psychology has yet to come to grips.     

Opposite influence of perceptual memory on initial and prolonged perception of sensory ambiguity

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.     

Characterization of the percepts evoked by discontinuous motion over the perioral skin.

The capacity of human subjects to process information about discontinuous and continuous movement was evaluated. Constant-velocity brushing stimuli were delivered through aperture plates that rested lightly upon the mandibular skin. Each plate consisted of either two spatially separated, slit-like openings or a single continuous, longer opening. It was discovered that percepts of smooth apparent motion were achieved with the split apertures (i.e., from discontinuous movement) for only limited ranges of stimulus velocity. Moreover, the optimal velocity supporting smooth apparent motion increased with the separation between the slit-like openings. In a second series of experiments, subjects' ability to discriminate opposing directions of discontinuous and continuous movement was evaluated. It was found that subjects could derive directional information from percepts elicited by discontinuous movement. However, the capacity to discriminate opposing directions of continuous movement cannot be explained solely in terms of the ability to process information about the change in position of a stimulus from its onset to its offset.     

CONTEXTUAL SHIFTS IN HEDONIC JUDGMENTS

Taste intensity ratings are higher when many low-intensity stimuli are presented than when many high-intensity stimuli are presented. It is investigated whether similar contextual shifts occur for hedonic judgments. Experiment 1 assessed the perceived pleasantness of a heterogeneous stimulus set containing either a large number of quinine (unpleasant context) or sucrose (pleasant context) solutions. In Experiment 2 subjects made hedonic judgments for a set of sucrose solutions. They were classified pre-experimentally as ‘sweet-likers,’‘neutrals,’ or ‘sweet-dislikers.’ Stimulus frequency distributions were positively or negatively skewed. In Experiment 1, hedonic ratings in the pleasant context were lower than those obtained in the unpleasant context. In Experiment 2, no effect of stimulus context was found. It is argued that contextual shifts in hedonic judgments are found only if the stimuli differ substantially along the hedonic dimension. In addition, the subject groups should be homogeneous with regard to preference rank orders, and should be sufficiently large to allow testing with adequate statistical power.     

Effects of the Ebbinghaus illusion on different behaviors: one- and two-handed grasping; one- and two-handed manual estimation; metric and comparative judgment

Many studies have suggested that visually-guided action is largely immune to the effects of several pictorial illusions that strongly influence perceptual judgments. The judgments in these experiments, however, have usually involved comparisons of multiple elements within a display, whereas the visually-guided actions have typically involved a pincer grip directed to only one display element. The three experiments presented here assess the influence of this confound on the perception versus action illusion dissociation. In general, the studies suggest (a) that the confound affects perceptual judgment but not grasping or manual estimation, and (b) that difficult visuomotor tasks are more affected by the Ebbinghaus illusion than easier tasks. In Experiment 1, participants reached for or made judgments about plastic disks placed in the center of the Ebbinghaus illusion display. Some participants reached for or made judgments about only the disk on the right, whereas others reached for or judged both disks simultaneously. A large effect of the illusion was found for grasping and comparative judgment, but not for manual estimation or metric judgment. In Experiment 2, the disks were elevated slightly to make gripping the targets easier, and the effects of the illusion on grasping were greatly reduced. For Experiment 3, participants performed the manual estimation task while the hands were placed in view, on the surface of the table, and the effects of the illusion were significantly increased. Taken together, the experiments indicate that task difficulty and hand visibility affect whether a task will be influenced by pictorial illusions or not. One- and two-handed grasping seem to be affected approximately equally.     

Network Connections That Evolve to Circumvent the Inverse Optics Problem

A fundamental problem in vision science is how useful perceptions and behaviors arise in the absence of information about the physical sources of retinal stimuli (the inverse optics problem). Psychophysical studies show that human observers contend with this problem by using the frequency of occurrence of stimulus patterns in cumulative experience to generate percepts. To begin to understand the neural mechanisms underlying this strategy, we examined the connectivity of simple neural networks evolved to respond according to the cumulative rank of stimulus luminance values. Evolved similarities with the connectivity of early level visual neurons suggests that biological visual circuitry uses the same mechanisms as a means of creating useful perceptions and behaviors without information about the real world.     

Symmetry detection across the visual field.

Humans are extremely sensitive to symmetry when it is foveated but sensitivity drops as a symmetrical region of a fixed size is moved into the periphery. A psychophysical study was undertaken to determine if eccentricity dependent sensitivity loss could be overcome by magnifying stimuli at each eccentricity (E) by a factor F = 1 + E/E2, where E2 indicates the eccentricity at which the size of a stimulus must be doubled, relative to a foveal standard, to achieve equivalent performance. The psychophysical task required subjects to decide on each trial in which of two intervals a symmetrical stimulus had been presented. Stimuli were presented at a range of sizes and eccentricities (0 to 8 degrees) and the probability of a correct discrimination was computed for each condition. In Experiment 1, thresholds were measured with stimuli set to maximum available contrast and, in Experiment 2, stimuli were presented at a constant multiple of contrast detection threshold. In both experiments, a single scaling function removed most of the eccentricity dependent variation from the data. However, the E2 value recovered for one subject tested in both experiments was larger by about 65% when stimuli were not equated for visibility. We conclude that symmetry detection can be equated across a range of eccentricities by scaling stimuli with an E2 in the range of 0.88 to 1.38 degrees. Failure to equate for visibility across all viewing conditions may result in an inflated estimate of E2.     

Detection of light intensity in the frog retina: evidence from dark and light adaptation

The frog retina was stimulated with light flashes homogeneous in space but not time. The time heterogeneity of stimulation was created by abrupt change of a referent stimulus for a stimulus with different luminance. Such changes form a time pattern, as well as sharp borders of luminance between the neighbor areas of the visual field form a spatial pattern. The electroretinogram recorded in response to presentation of a triad of stimuli: the onset of a short flash of homogeneous light after long dark (or light) adaptation of a retina, brief sequence of the referent and test light flashes varied in luminance, and the offset, with returning to the initial level of adaptation. It was shown that responses of the retina under conditions of time heterogeneity of stimulation could be divided in two types as well as under conditions of spatial heterogeneity. Such a dual change in amplitude confirms our earlier hypothesis on the existence of two mechanisms of luminance coding in the frog retina. The first mechanism encodes power characteristics of light, it forms the information on the absolute level of the environmental luminance. Its activity is connected basically with receptors and cells of the external plexiform layer of the frog's retina. It is responsible for the b-wave of the electroretinogram. The other mechanism associated with RERG is based on a vector code of stimuli. This mechanism forms the information on spatial and time differentiation of the light flow in the visual field and is connected basically with cells of the internal plexiform layer. The results suggest that the frog retina has the individual mechanism for time pattern detection, distinguishing it from the homogeneous light flow in a similar way as in case of spatial light pattern detection. It is possible that the first mechanism is responsible for the detection of any new stimulus in general, irrespective of its specificity, whereas the second mechanism serves for the measurement of suprathreshold differences between stimuli.     

A neural network model for stroboscopic alternative motion

A neural network which models multistable perception is presented. The network consists of sensor and inner neurons. The dynamics is established by a stochastic neuronal dynamics, a formal Hebb-type coupling dynamics and a resource mechanism that corresponds to saturation effects in perception. From this a system of coupled differential equations is derived and analyzed. Single stimuli are bound to exactly one percept, even in ambiguous situations where multistability occurs. The network exhibits discontinuous as well as continuous phase transitions and models various empirical findings, including the percepts of succession, alternative motion and simultaneity; the percept of oscillation is explained by oscillating percepts at a continuous phase transition. Received: 13 September 1995 / Accepted: 3 June 1996     

Binocular onset rivalry at the time of saccades and stimulus jumps

Recent studies suggest that binocular rivalry at stimulus onset, so called onset rivalry, differs from rivalry during sustained viewing. These observations raise the interesting question whether there is a relation between onset rivalry and rivalry in the presence of eye movements. We therefore studied binocular rivalry when stimuli jumped from one visual hemifield to the other, either through a saccade or through a passive stimulus displacement, and we compared rivalry after such displacements with onset and sustained rivalry. We presented opponent motion, orthogonal gratings and face/house stimuli through a stereoscope. For all three stimulus types we found that subjects showed a strong preference for stimuli in one eye or one hemifield (Experiment 1), and that these subject-specific biases did not persist during sustained viewing (Experiment 2). These results confirm and extend previous findings obtained with gratings. The results from the main experiment (Experiment 3) showed that after a passive stimulus jump, switching probability was low when the preferred eye was dominant before a stimulus jump, but when the non-preferred eye was dominant beforehand, switching probability was comparatively high. The results thus showed that dominance after a stimulus jump was tightly related to eye dominance at stimulus onset. In the saccade condition, however, these subject-specific biases were systematically reduced, indicating that the influence of saccades can be understood from a systematic attenuation of the subjects' onset rivalry biases. Taken together, our findings demonstrate a relation between onset rivalry and rivalry after retinal shifts and involvement of extra-retinal signals in binocular rivalry.