Crowding by invisible flankers

Background

Human object recognition degrades sharply as the target object moves from central vision into peripheral vision. In particular, one''s ability to recognize a peripheral target is severely impaired by the presence of flanking objects, a phenomenon known as visual crowding. Recent studies on how visual awareness of flanker existence influences crowding had shown mixed results. More importantly, it is not known whether conscious awareness of the existence of both the target and flankers are necessary for crowding to occur.

Methodology/Principal Findings

Here we show that crowding persists even when people are completely unaware of the flankers, which are rendered invisible through the continuous flash suppression technique. Contrast threshold for identifying the orientation of a grating pattern was elevated in the flanked condition, even when the subjects reported that they were unaware of the perceptually suppressed flankers. Moreover, we find that orientation-specific adaptation is attenuated by flankers even when both the target and flankers are invisible.

Conclusions

These findings complement the suggested correlation between crowding and visual awareness. What''s more, our results demonstrate that conscious awareness and attention are not prerequisite for crowding.     

Grouping and Crowding Affect Target Appearance over Different Spatial Scales

Crowding is the impairment of peripheral target perception by nearby flankers. A number of recent studies have shown that crowding shares many features with grouping. Here, we investigate whether effects of crowding and grouping on target perception are related by asking whether they operate over the same spatial scale. A target letter T had two sets of flanking Ts of varying orientations. The first set was presented close to the target, yielding strong crowding. The second set was either close enough to cause crowding on their own or too far to cause crowding on their own. The Ts of the second set had the same orientation that either matched the target’s orientation (Grouped condition) or not (Ungrouped condition). In Experiment 1, the Grouped flankers reduced crowding independently of their distance from the target, suggesting that grouping operated over larger distances than crowding. In Experiments 2 and 3 we found that grouping did not affect sensitivity but produced a strong bias to report that the grouped orientation was present at the target location whether or not it was. Finally, we investigated whether this bias was a response or perceptual bias, rejecting the former in favor of a perceptual grouping explanation. We suggest that the effect of grouping is to assimilate the target to the identity of surrounding flankers when they are all the same, and that this shape assimilation effect differs in its spatial scale from the integration effect of crowding.     

Reducing crowding by weakening inhibitory lateral interactions in the periphery with perceptual learning

We investigated whether lateral masking in the near-periphery, due to inhibitory lateral interactions at an early level of central visual processing, could be weakened by perceptual learning and whether learning transferred to an untrained, higher-level lateral masking known as crowding. The trained task was contrast detection of a Gabor target presented in the near periphery (4°) in the presence of co-oriented and co-aligned high contrast Gabor flankers, which featured different target-to-flankers separations along the vertical axis that varied from 2λ to 8λ. We found both suppressive and facilitatory lateral interactions at target-to-flankers distances (2λ - 4λ and 8λ, respectively) that were larger than those found in the fovea. Training reduces suppression but does not increase facilitation. Most importantly, we found that learning reduces crowding and improves contrast sensitivity, but has no effect on visual acuity (VA). These results suggest a different pattern of connectivity in the periphery with respect to the fovea as well as a different modulation of this connectivity via perceptual learning that not only reduces low-level lateral masking but also reduces crowding. These results have important implications for the rehabilitation of low-vision patients who must use peripheral vision to perform tasks, such as reading and refined figure-ground segmentation, which normal sighted subjects perform in the fovea.     

Distractor-target interactions during directed visual attention

The spatial extent of directed visual attention (DVA) was examined in a series of experiments using precuing in a suprathreshold luminance detection (reaction time) paradigm. Previous findings (Hughes, H. C. and Zimba, L. D. J. Exp. Psychol.; Human Percept Perf., 1985, 11, 409-430) indicated that, in an empty visual field, the effects of DVA were primarily manifest as a uniform elevation of response times to all probe targets in the hemifield contralateral to the observer's expectancy. The present experiments were designed to determine whether increased spatial selectivity could be found when luminous markers indicated the exact location of the expected visual target. To maintain equivalent states of adaptation in both hemifields, luminous markers were also present at the same location in the contralateral hemifield. In general, hemifield effects were again obtained, but with two notable exceptions. First, marking locations in the unattended hemifield produced a local increase (enhanced interference) in RTs above the level characteristic of other locations within that hemifield. Second, when multiple locations were indicated with identical luminous markers, graded costs were obtained in both hemifields. However, scaling the markers according to estimates of cortical magnification factor (M) substantially reduced the slope of these inhibitory gradients, and the results once again approached those characteristic of an unstructured visual field. The findings suggest that when attention is directed to a marked location along the horizontal meridian, a transition in performance typically occurs at the vertical meridian. In addition, irrelevant stimuli some distance from the attentional focus interfere with detection times to unexpected targets that appear in the same vicinity. This interference may relate to an enhanced susceptibility to spatial interactions between the distractors and target away from the attentional focus. The interference appears to extend over a constant area of visual cortex, since it is reduced when the markers are M-scaled.     

Top-down modulation of lateral interactions in early vision: does attention affect integration of the whole or just perception of the parts?

Attention can modulate sensitivity to local stimuli in early vision. But, can attention also modulate integration of local stimuli into global visual patterns? We recently measured effects of attention on the phenomenon of lateral interactions between collinear elements, commonly thought to reflect long-range mechanisms in early visual cortex underlying contour integration. We showed improved detection of low-contrast central Gabor targets in the context of collinear flankers, but only when the collinear flankers were attended for a secondary task rather than ignored in favor of an orthogonal flanker pair. Here, we contrast two hypotheses for how attention might modulate flanker influences on the target: by changing just local sensitivity to the flankers themselves (flanker-modulation-only hypothesis), or by weighting integrative connections between flanker and target (connection-weighting hypothesis). Modeled on the known nonlinear dependence of target visibility on collinear flanker contrast, the first hypothesis predicts that an increase in physical flanker contrast should readily offset any reduction in their effective contrast when ignored, thus eliminating attentional modulation. Conversely, the second hypothesis predicts that attentional modulation should persist even for the highest flanker contrasts. Our results showed the latter outcome and indicated that attention modulates flanker-target integration, rather than just processing of local flanker elements.     

Visual search for a target changing in synchrony with an auditory signal

We examined whether the detection of audio-visual temporal synchrony is determined by a pre-attentive parallel process, or by an attentive serial process using a visual search paradigm. We found that detection of a visual target that changed in synchrony with an auditory stimulus was gradually impaired as the number of unsynchronized visual distractors increased (experiment 1), whereas synchrony discrimination of an attended target in a pre-cued location was unaffected by the presence of distractors (experiment 2). The effect of distractors cannot be ascribed to reduced target visibility nor can the increase in false alarm rates be predicted by a noisy parallel processing model. Reaction times for target detection increased linearly with number of distractors, with the slope being about twice as steep for target-absent trials as for target-present trials (experiment 3). Similar results were obtained regardless of whether the audio-visual stimulus consisted of visual flashes synchronized with amplitude-modulated pips, or of visual rotations synchronized with frequency-modulated up-down sweeps. All of the results indicate that audio-visual perceptual synchrony is judged by a serial process and are consistent with the suggestion that audio-visual temporal synchrony is detected by a 'mid-level' feature matching process.     

Shrinking Bouma’s window: How to model crowding in dense displays

In crowding, perception of a target deteriorates in the presence of nearby flankers. Traditionally, it is thought that visual crowding obeys Bouma’s law, i.e., all elements within a certain distance interfere with the target, and that adding more elements always leads to stronger crowding. Crowding is predominantly studied using sparse displays (a target surrounded by a few flankers). However, many studies have shown that this approach leads to wrong conclusions about human vision. Van der Burg and colleagues proposed a paradigm to measure crowding in dense displays using genetic algorithms. Displays were selected and combined over several generations to maximize human performance. In contrast to Bouma’s law, only the target’s nearest neighbours affected performance. Here, we tested various models to explain these results. We used the same genetic algorithm, but instead of selecting displays based on human performance we selected displays based on the model’s outputs. We found that all models based on the traditional feedforward pooling framework of vision were unable to reproduce human behaviour. In contrast, all models involving a dedicated grouping stage explained the results successfully. We show how traditional models can be improved by adding a grouping stage.     

Visual crowding is correlated with awareness

Crowding by nearby features causes identification failures in the peripheral visual field. However, prominent visual features can sometimes fail to reach awareness, causing scenes to be incorrectly interpreted. Here we examine whether awareness of the flanking features is necessary for crowding to occur. Flankers that were physically present were rendered perceptually absent with adaptation-induced blindness. In a letter identification task, targets were presented unflanked or with up to four flanker letters. On each trial, observers reported both the number of letters they perceived and the identity of a target letter. This paradigm allowed trial-by-trial assessment of awareness and crowding and ensured that both targets and flankers were attended. Target-letter identification performance was correlated with the number of flanking letters that were perceived on a given trial, regardless of the number that were physically present. Our data demonstrate that crowding can be released when flanking elements at attended locations are suppressed from visual awareness.     

Attentional modulation of temporal contrast sensitivity in human vision

Recent psychophysical studies have shown that attention can alter contrast sensitivities for temporally broadband stimuli such as flashed gratings. The present study examined the effect of attention on the contrast sensitivity for temporally narrowband stimuli with various temporal frequencies. Observers were asked to detect a drifting grating of 0-40 Hz presented gradually in the peripheral visual field with or without a concurrent letter identification task in the fovea. We found that removal of attention by the concurrent task reduced the contrast sensitivity for gratings with low temporal frequencies much more profoundly than for gratings with high temporal frequencies and for flashed gratings. The analysis revealed that the temporal contrast sensitivity function had a more band-pass shape with poor attention. Additional experiments showed that this was also true when the target was presented in various levels of luminance noise. These results suggest that regardless of the presence of external noise, attention extensively modulates visual sensitivity for sustained retinal inputs.     

Touch Influences Visual Perception with a Tight Orientation-Tuning

Stimuli from different sensory modalities are thought to be processed initially in distinct unisensory brain areas prior to convergence in multisensory areas. However, signals in one modality can influence the processing of signals from other modalities and recent studies suggest this cross-modal influence may occur early on, even in ‘unisensory’ areas. Some recent psychophysical studies have shown specific cross-modal effects between touch and vision during binocular rivalry, but these cannot completely rule out a response bias. To test for genuine cross-modal integration of haptic and visual signals, we investigated whether congruent haptic input could influence visual contrast sensitivity compared to incongruent haptic input in three psychophysical experiments using a two-interval, two-alternative forced-choice method to eliminate response bias. The initial experiment demonstrated that contrast thresholds for a visual grating were lower when exploring a haptic grating that shared the same orientation compared to an orthogonal orientation. Two subsequent experiments mapped the orientation and spatial frequency tunings for the congruent haptic facilitation of vision, finding a clear orientation tuning effect but not a spatial frequency tuning. In addition to an increased contrast sensitivity for iso-oriented visual-haptic gratings, we found a significant loss of sensitivity for orthogonally oriented visual-haptic gratings. We conclude that the tactile influence on vision is a result of a tactile input to orientation-tuned visual areas.     

Visual Benefits in Apparent Motion Displays: Automatically Driven Spatial and Temporal Anticipation Are Partially Dissociated

Many behaviourally relevant sensory events such as motion stimuli and speech have an intrinsic spatio-temporal structure. This will engage intentional and most likely unintentional (automatic) prediction mechanisms enhancing the perception of upcoming stimuli in the event stream. Here we sought to probe the anticipatory processes that are automatically driven by rhythmic input streams in terms of their spatial and temporal components. To this end, we employed an apparent visual motion paradigm testing the effects of pre-target motion on lateralized visual target discrimination. The motion stimuli either moved towards or away from peripheral target positions (valid vs. invalid spatial motion cueing) at a rhythmic or arrhythmic pace (valid vs. invalid temporal motion cueing). Crucially, we emphasized automatic motion-induced anticipatory processes by rendering the motion stimuli non-predictive of upcoming target position (by design) and task-irrelevant (by instruction), and by creating instead endogenous (orthogonal) expectations using symbolic cueing. Our data revealed that the apparent motion cues automatically engaged both spatial and temporal anticipatory processes, but that these processes were dissociated. We further found evidence for lateralisation of anticipatory temporal but not spatial processes. This indicates that distinct mechanisms may drive automatic spatial and temporal extrapolation of upcoming events from rhythmic event streams. This contrasts with previous findings that instead suggest an interaction between spatial and temporal attention processes when endogenously driven. Our results further highlight the need for isolating intentional from unintentional processes for better understanding the various anticipatory mechanisms engaged in processing behaviourally relevant stimuli with predictable spatio-temporal structure such as motion and speech.     

Size as a parameter for tuning visual attention]

The ability of visual attention to tune to the stimulus size (when this size could not be described by spatial frequencies) was studies. Sinusoidal gratings with frequencies of 1.5, 3, and 6 cycle/degree were used as test stimuli. All these stimuli consisted of 3 periods, consequently, they had different sizes: 2 x 2, 1 x 1, and 0.5 x 0.5 degrees. Three reference stimuli had the same sizes but were constructed as a superposition of all the test frequencies. The reference stimulus of suprathreshold contrast was displayed for 400 ms to the left or to the right of a fixation point at a distance of 3 degrees. After that, the test stimulus of threshold contrast was for 100 ms displayed symmetrically to the fixation point on the other side. Subjects were instructed that the sizes of the reference and test stimuli were the same. It was found that the probability of test detection decreased with increase in the difference between the sizes of the reference and test stimuli. Since in our experiments the spatial frequency could not be used for tuning visual attention, the obtained results suggest that there are specialized mechanisms in the visual system for estimation of the general image size.     

Effects of Spatial Frequency Similarity and Dissimilarity on Contour Integration

We examined the effects of spatial frequency similarity and dissimilarity on human contour integration under various conditions of uncertainty. Participants performed a temporal 2AFC contour detection task. Spatial frequency jitter up to 3.0 octaves was applied either to background elements, or to contour and background elements, or to none of both. Results converge on four major findings. (1) Contours defined by spatial frequency similarity alone are only scarcely visible, suggesting the absence of specialized cortical routines for shape detection based on spatial frequency similarity. (2) When orientation collinearity and spatial frequency similarity are combined along a contour, performance amplifies far beyond probability summation when compared to the fully heterogenous condition but only to a margin compatible with probability summation when compared to the fully homogenous case. (3) Psychometric functions are steeper but not shifted for homogenous contours in heterogenous backgrounds indicating an advantageous signal-to-noise ratio. The additional similarity cue therefore not so much improves contour detection performance but primarily reduces observer uncertainty about whether a potential candidate is a contour or just a false positive. (4) Contour integration is a broadband mechanism which is only moderately impaired by spatial frequency dissimilarity.     

Colour Terms Affect Detection of Colour and Colour-Associated Objects Suppressed from Visual Awareness

The idea that language can affect how we see the world continues to create controversy. A potentially important study in this field has shown that when an object is suppressed from visual awareness using continuous flash suppression (a form of binocular rivalry), detection of the object is differently affected by a preceding word prime depending on whether the prime matches or does not match the object. This may suggest that language can affect early stages of vision. We replicated this paradigm and further investigated whether colour terms likewise influence the detection of colours or colour-associated object images suppressed from visual awareness by continuous flash suppression. This method presents rapidly changing visual noise to one eye while the target stimulus is presented to the other. It has been shown to delay conscious perception of a target for up to several minutes. In Experiment 1 we presented greyscale photos of objects. They were either preceded by a congruent object label, an incongruent label, or white noise. Detection sensitivity (d’) and hit rates were significantly poorer for suppressed objects preceded by an incongruent label compared to a congruent label or noise. In Experiment 2, targets were coloured discs preceded by a colour term. Detection sensitivity was significantly worse for suppressed colour patches preceded by an incongruent colour term as compared to a congruent term or white noise. In Experiment 3 targets were suppressed greyscale object images preceded by an auditory presentation of a colour term. On congruent trials the colour term matched the object’s stereotypical colour and on incongruent trials the colour term mismatched. Detection sensitivity was significantly poorer on incongruent trials than congruent trials. Overall, these findings suggest that colour terms affect awareness of coloured stimuli and colour- associated objects, and provide new evidence for language-perception interaction in the brain.     

Realigning Thunder and Lightning: Temporal Adaptation to Spatiotemporally Distant Events

The brain is able to realign asynchronous signals that approximately coincide in both space and time. Given that many experience-based links between visual and auditory stimuli are established in the absence of spatiotemporal proximity, we investigated whether or not temporal realignment arises in these conditions. Participants received a 3-min exposure to visual and auditory stimuli that were separated by 706 ms and appeared either from the same (Experiment 1) or from different spatial positions (Experiment 2). A simultaneity judgment task (SJ) was administered right afterwards. Temporal realignment between vision and audition was observed, in both Experiment 1 and 2, when comparing the participants’ SJs after this exposure phase with those obtained after a baseline exposure to audiovisual synchrony. However, this effect was present only when the visual stimuli preceded the auditory stimuli during the exposure to asynchrony. A similar pattern of results (temporal realignment after exposure to visual-leading asynchrony but not after exposure to auditory-leading asynchrony) was obtained using temporal order judgments (TOJs) instead of SJs (Experiment 3). Taken together, these results suggest that temporal recalibration still occurs for visual and auditory stimuli that fall clearly outside the so-called temporal window for multisensory integration and appear from different spatial positions. This temporal realignment may be modulated by long-term experience with the kind of asynchrony (vision-leading) that we most frequently encounter in the outside world (e.g., while perceiving distant events).     

Eye position affects orienting of visuospatial attention

The ability to detect an incoming visual stimulus is enhanced by knowledge of stimulus location (orienting of visuospatial attention). Although the brain mechanisms at the basis of this enhancement are not yet fully clarified, there is evidence that orienting of attention is accompanied by the activation of oculomotor circuits. It remains unclear, however, whether this oculomotor activity is an epiphenomenon or is functionally related to the attentional process. Attentional benefits are usually measured by the classical Posner paradigm. When subjects fixate centrally and are requested to detect a visual stimulus that could appear in an attended or unattended location, they react faster to stimuli appearing in the attended one. Here, we demonstrate that in monocular vision visuospatial attention was significantly modulated by the position of the eye in the orbit. When the screen was placed 40 degrees to the right or to the left of subjects' sagittal plane, attentional benefits for stimuli appearing in subjects' temporal spatial hemifield dramatically decayed, even if the retinal stimulation was exactly the same as in the classical paradigm. The finding that eyes and attention show a common limit stop point supports their close functional coupling.     

Negative priming under rapid serial visual presentation

Negative priming (NP) was examined under a new paradigm wherein a target and distractors were temporally separated using rapid serial visual presentation (RSVP). The results from the two experiments revealed that (a) NP was robust under RSVP, such that the responses to a target were slower when the target served as a distractor in a previous trial than when it did not; (b) NP was found regardless of whether the distractors appeared before or after the targets; and (c) NP was stronger when the distractor was more distinctive. These findings are generally similar to those on NP in the spatial search task. The implications for the processes causing NP under RSVP are discussed in the current paper.     

A first- and second-order motion energy analysis of peripheral motion illusions leads to further evidence of "feature blur" in peripheral vision

Background

Anatomical and physiological differences between the central and peripheral visual systems are well documented. Recent findings have suggested that vision in the periphery is not just a scaled version of foveal vision, but rather is relatively poor at representing spatial and temporal phase and other visual features. Shapiro, Lu, Huang, Knight, and Ennis (2010) have recently examined a motion stimulus (the “curveball illusion”) in which the shift from foveal to peripheral viewing results in a dramatic spatial/temporal discontinuity. Here, we apply a similar analysis to a range of other spatial/temporal configurations that create perceptual conflict between foveal and peripheral vision.

Methodology/Principal Findings

To elucidate how the differences between foveal and peripheral vision affect super-threshold vision, we created a series of complex visual displays that contain opposing sources of motion information. The displays (referred to as the peripheral escalator illusion, peripheral acceleration and deceleration illusions, rotating reversals illusion, and disappearing squares illusion) create dramatically different perceptions when viewed foveally versus peripherally. We compute the first-order and second-order directional motion energy available in the displays using a three-dimensional Fourier analysis in the (x, y, t) space. The peripheral escalator, acceleration and deceleration illusions and rotating reversals illusion all show a similar trend: in the fovea, the first-order motion energy and second-order motion energy can be perceptually separated from each other; in the periphery, the perception seems to correspond to a combination of the multiple sources of motion information. The disappearing squares illusion shows that the ability to assemble the features of Kanisza squares becomes slower in the periphery.

Conclusions/Significance

The results lead us to hypothesize “feature blur” in the periphery (i.e., the peripheral visual system combines features that the foveal visual system can separate). Feature blur is of general importance because humans are frequently bringing the information in the periphery to the fovea and vice versa.     

Visual advantage in deaf adults linked to retinal changes

The altered sensory experience of profound early onset deafness provokes sometimes large scale neural reorganisations. In particular, auditory-visual cross-modal plasticity occurs, wherein redundant auditory cortex becomes recruited to vision. However, the effect of human deafness on neural structures involved in visual processing prior to the visual cortex has never been investigated, either in humans or animals. We investigated neural changes at the retina and optic nerve head in profoundly deaf (N = 14) and hearing (N = 15) adults using Optical Coherence Tomography (OCT), an in-vivo light interference method of quantifying retinal micro-structure. We compared retinal changes with behavioural results from the same deaf and hearing adults, measuring sensitivity in the peripheral visual field using Goldmann perimetry. Deaf adults had significantly larger neural rim areas, within the optic nerve head in comparison to hearing controls suggesting greater retinal ganglion cell number. Deaf adults also demonstrated significantly larger visual field areas (indicating greater peripheral sensitivity) than controls. Furthermore, neural rim area was significantly correlated with visual field area in both deaf and hearing adults. Deaf adults also showed a significantly different pattern of retinal nerve fibre layer (RNFL) distribution compared to controls. Significant correlations between the depth of the RNFL at the inferior-nasal peripapillary retina and the corresponding far temporal and superior temporal visual field areas (sensitivity) were found. Our results show that cross-modal plasticity after early onset deafness may not be limited to the sensory cortices, noting specific retinal adaptations in early onset deaf adults which are significantly correlated with peripheral vision sensitivity.     

Spatiotopic visual maps revealed by saccadic adaptation in humans

Saccadic adaptation [1] is a powerful experimental paradigm to probe the mechanisms of eye movement control and spatial vision, in which saccadic amplitudes change in response to false visual feedback. The adaptation occurs primarily in the motor system [2, 3], but there is also evidence for visual adaptation, depending on the size and the permanence of the postsaccadic error [4-7]. Here we confirm that adaptation has a strong visual component and show that the visual component of the adaptation is spatially selective in external, not retinal coordinates. Subjects performed?a memory-guided, double-saccade, outward-adaptation task designed to maximize visual adaptation and to dissociate the visual and motor corrections. When the memorized saccadic target was in the same position (in external space) as that used in the adaptation training, saccade targeting was strongly influenced by adaptation (even if not matched in retinal or cranial position), but when in the same retinal or cranial but different external spatial position, targeting was unaffected by adaptation, demonstrating unequivocal spatiotopic selectivity. These results point to the existence of a spatiotopic neural representation for eye movement control that adapts in response to saccade error signals.