Visual Consciousness Research Articles

Roles of contour and surface processing in microgenesis of object perception and visual consciousness.

Developments in visual neuroscience and neural-network modeling indicate the existence of separate pathways for the processing of form and surface attributes of a visual object. In line with prior theoretical proposals, it is assumed that the processing of form can be explicit or conscious only as or after the surface property such as color is filled in. In conjunction with extant psychophysical findings, these developments point to interesting distinctions between nonconscious and conscious processing of these attributes, specifically in relation to distinguishable temporal dynamics. At nonconscious levels form processing proceeds faster than surface processing, whereas in contrast, at conscious levels form processing proceeds slower than surface processing. I mplications of separate form and surface processing for current and future psychophysical and neuroscientific research, particularly that relating cortical oscillations to conjunctions of surface and form features, and for cognitive science and philosophy of mind and consciousness are discussed.

Visual Processing in Rapid-Chase Systems: Image Processing, Attention, and Awareness

Visual stimuli can be classified so rapidly that their analysis may be based on a single sweep of feedforward processing through the visuomotor system. Behavioral criteria for feedforward processing can be evaluated in response priming tasks where speeded pointing or keypress responses are performed toward target stimuli which are preceded by prime stimuli. We apply this method to several classes of complex stimuli. (1) When participants classify natural images into animals or non-animals, the time course of their pointing responses indicates that prime and target signals remain strictly sequential throughout all processing stages, meeting stringent behavioral criteria for feedforward processing (rapid-chase criteria). (2) Such priming effects are boosted by selective visual attention for positions, shapes, and colors, in a way consistent with bottom-up enhancement of visuomotor processing, even when primes cannot be consciously identified. (3) Speeded processing of phobic images is observed in participants specifically fearful of spiders or snakes, suggesting enhancement of feedforward processing by long-term perceptual learning. (4) When the perceived brightness of primes in complex displays is altered by means of illumination or transparency illusions, priming effects in speeded keypress responses can systematically contradict subjective brightness judgments, such that one prime appears brighter than the other but activates motor responses as if it was darker. We propose that response priming captures the output of the first feedforward pass of visual signals through the visuomotor system, and that this output lacks some characteristic features of more elaborate, recurrent processing. This way, visuomotor measures may become dissociated from several aspects of conscious vision. We argue that “fast” visuomotor measures predominantly driven by feedforward processing should supplement “slow” psychophysical measures predominantly based on visual awareness.

Are there unconscious perceptual processes?

Blindsight and vision for action seem to be exemplars of unconscious visual processes. However, researchers have recently argued that blindsight is not really a kind of unconscious vision but is rather severely degraded conscious vision. Morten Overgaard and colleagues have recently developed new methods for measuring the visibility of visual stimuli. Studies using these methods show that reported clarity of visual stimuli correlates with accuracy in both normal individuals and blindsight patients. Vision for action has also come under scrutiny. Recent findings seem to show that information processed by the dorsal stream for online action contributes to visual awareness. Some interpret these results as showing that some dorsal stream processes are conscious visual processes (e.g., Gallese, 2007; Jacob & Jeannerod, 2003). The aim of this paper is to provide new support for the more traditional view that blindsight and vision for action are genuinely unconscious perceptual processes. I argue that individuals with blindsight do not have access to the kind of purely qualitative color and size information which normal individuals do. So, even though people with blindsight have a kind of cognitive consciousness, visual information processing in blindsight patients is not associated with a distinctly visual phenomenology. I argue further that while dorsal stream processing seems to contribute to visual awareness, only information processed by the early dorsal stream (V1, V2, and V3) is broadcast to working memory. Information processed by later parts of the dorsal stream (the parietal lobe) never reaches working memory and hence does not correlate with phenomenal awareness. I conclude that both blindsight and vision for action are genuinely unconscious visual processes.

The primary visual cortex, and feedback to it, are not necessary for conscious vision

A compelling single case report of visual awareness (visual qualia) without primary visual cortex would be sufficient to refute the hypothesis that the primary visual cortex and the back-projections to it are necessary for conscious visual experience. In a previous study, we emphasized the presence of crude visual awareness in Patient G.Y., with a lesion of the primary visual cortex, who is aware of, and able to discriminate, fast-moving visual stimuli presented to his blind field. The visual nature of Patient G.Y.'s blind field experience has since been questioned and it has been suggested that the special circumstances of repeated testing over decades may have altered Patient G.Y.'s visual pathways. We therefore sought new evidence of visual awareness without primary visual cortex in patients for whom such considerations do not apply. Three patients with hemianopic field defects (Patient G.N. and Patient F.B. with MRI confirmed primary visual cortex lesions, Patient C.G. with an inferred lesion) underwent detailed psychophysical testing in their blind fields. Visual stimuli were presented at different velocities and contrasts in two- and four-direction discrimination experiments and the direction of motion and awareness reported using a forced-choice paradigm. Detailed verbal reports were also obtained of the nature of the blind field experience with comparison of the drawings of the stimulus presented in the blind and intact fields, where possible. All three patients reported visual awareness in their blind fields. Visual awareness was significantly more likely when a moving stimulus was present compared to no stimulus catch trials (P < 0.01 for each subject). Psychophysical performance in Patient F.B. and Patient G.N. was consistent with the Riddoch syndrome, with higher levels of visual awareness for moving compared to static stimuli (P < 0.001) and intact direction discrimination (P < 0.0001 for two- and four-direction experiments). Although the blind field experience of all three subjects was degraded, it was clearly visual in nature. We conclude that the primary visual cortex or back-projections to it are not necessary for visual awareness.

Influence of Near Threshold Visual Distractors on Perceptual Detection and Reaching Movements

Providing evidence against a dissociation between conscious vision for perception and unconscious vision for action, recent studies have suggested that perceptual and motor decisions are based on a unique signal but distinct decisional thresholds. The aim of the present study was to provide a direct test of this assumption in a perceptual-motor dual task involving arm movements. In 300 trials, 10 participants performed speeded pointing movements toward a highly visible target located at 10° from the fixation point and ± 45° from the body midline. The target was preceded by one or two close to threshold distractor(s) (80 ms stimulus onset asynchrony) presented ± 30° according to the target location. After each pointing movement, participants judged whether the distractor was present or not on either side of the target. Results showed a robust reaction time facilitation effect and a deviation toward the distractor when the distractor was both present and consciously perceived (Hit). A small reaction time facilitation was also observed when two distractors were physically present but undetected (double-miss)--this facilitation being highly correlated with the physical contrast of the distractors. These results are compatible with the theory proposing that perceptual and motor decisions are based on a common signal but emerge from a contrast dependent fixed threshold for motor responses and a variable context dependent criterion for perceptual responses. This paper thus extends to arm movement control previous findings related to oculomotor control.

Response priming driven by local contrast, not subjective brightness

We demonstrate qualitative dissociations of brightness processing in visuomotor priming and conscious vision. Speeded keypress responses to the brighter of two luminance targets were performed in the presence of preceding dark and bright primes (clearly visible and flanking the targets) whose apparent brightness values were enhanced or attenuated by a visual illusion. Response times to the targets were greatly affected by consistent versus inconsistent arrangements of the primes, relative to the targets (response priming). Priming effects could systematically contradict subjective brightness matches, such that one prime could appear brighter than the other but could prime as if it were darker. Systematic variation of the illusion showed that response-priming effects depended only on local flanker-background contrast, not on the subjective appearance of the flankers. Our findings suggest that speeded motor responses, as opposed to conscious perceptual judgments, access an early phase of lightness and brightness processing prior to full lightness constancy.

Response priming driven by local contrast, not subjective brightness

We demonstrate qualitative dissociations of brightness processing in visuomotor priming and conscious vision. Speeded keypress responses to the brighter of two luminance targets were performed in the presence of preceding dark and bright primes (clearly visible and flanking the targets) whose apparent brightness values were enhanced or attenuated by a visual illusion. Response times to the targets were greatly affected by consistent versus inconsistent arrangements of the primes, relative to the targets (response priming). Priming effects could systematically contradict subjective brightness matches, such that one prime could appear brighter than the other but could prime as if it were darker. Systematic variation of the illusion showed that response-priming effects depended only on local flanker-background contrast, not on the subjective appearance of the flankers. Our findings suggest that speeded motor responses, as opposed to conscious perceptual judgments, access an early phase of lightness and brightness processing prior to full lightness constancy.

Two Phases of V1 Activity for Visual Recognition of Natural Images

Present theories of visual recognition emphasize the role of interactive processing across populations of neurons within a given network, but the nature of these interactions remains unresolved. In particular, data describing the sufficiency of feedforward algorithms for conscious vision and studies revealing the functional relevance of feedback connections to the striate cortex seem to offer contradictory accounts of visual information processing. TMS is a good method to experimentally address this issue, given its excellent temporal resolution and its capacity to establish causal relations between brain function and behavior. We studied 20 healthy volunteers in a visual recognition task. Subjects were briefly presented with images of animals (birds or mammals) in natural scenes and were asked to indicate the animal category. MRI-guided stereotaxic single TMS pulses were used to transiently disrupt striate cortex function at different times after image onset (SOA). Visual recognition was significantly impaired when TMS was applied over the occipital pole at SOAs of 100 and 220 msec. The first interval has consistently been described in previous TMS studies and is explained as the interruption of the feedforward volley of activity. Given the late latency and discrete nature of the second peak, we hypothesize that it represents the disruption of a feedback projection to V1, probably from other areas in the visual network. These results provide causal evidence for the necessity of recurrent interactive processing, through feedforward and feedback connections, in visual recognition of natural complex images.

Magyar Szemleand the conservative mobilization againstVölkischideology and GermanVolksgeschichtein 1930s Hungary

In the 1930s in Hungary, the periodical Magyar Szemle (Hungarian Review) ranked as the foremost intellectual review of conservative thinking. Edited by the pro-establishment historian Gyula Szekfu, the journal provided important intellectual ammunition to the traditionalists of the right, in other words those who for various reasons sought to hold on to Istvan Bethlen’s version of moderate conservatism in ideology and a parliamentary system of limited pluralism and authoritarian checks in practice. The 1930s, however, bore witness to several challenges to the Horthy regime. The rise of the extreme right and the emancipatory (though often also fervently nationalist) program of the so-called nepi (populist or narodniki) writers presented coherent political alternatives to the prevailing order for the first time since the marginalization and emaciation of the left in the wake of the 1918–19 revolutions. Simultaneously, the country had to grapple with the emergence of Nazi Germany as an expansionist great power in the region. In this complicated situation, authors of Magyar Szemle confronted what they perceived as a dual threat: the increasing appeal of German imperialism and German political and historical thinking. Many intellectuals of the time, feeling that the German political challenge should be resisted through the adoption and adaptation of innovative German thinking on politics and history, espoused the new ideologies emanating from the unquestioned cultural center of Central Europe in some form. Magyar Szemle, however, emerged as a hub for public intellectuals who sought to hold on to a conservatism both more traditional and more open to some of the ideas of liberalism and who refused to abandon the established view of Hungarian history for a more ethnically conscious vision of the past. In the context of the dual German challenge of the 1930s, Magyar Szemle represented a site of intellectual resistance not so much against direct German political ambitions but against the new wave of German political thought and interpretations of history.

Opposing effects of attention and consciousness on afterimages

The brain's ability to handle sensory information is influenced by both selective attention and consciousness. There is no consensus on the exact relationship between these two processes and whether they are distinct. So far, no experiment has simultaneously manipulated both. We carried out a full factorial 2 x 2 study of the simultaneous influences of attention and consciousness (as assayed by visibility) on perception, correcting for possible concurrent changes in attention and consciousness. We investigated the duration of afterimages for all four combinations of high versus low attention and visible versus invisible. We show that selective attention and visual consciousness have opposite effects: paying attention to the grating decreases the duration of its afterimage, whereas consciously seeing the grating increases the afterimage duration. These findings provide clear evidence for distinctive influences of selective attention and consciousness on visual perception.

Backward Masking and Unmasking Across Saccadic Eye Movements

Humans make several eye movements every second, and thus a fundamental challenge in conscious vision is to maintain continuity by matching object representations in constantly shifting retinal coordinates. One possible mechanism for visual stability is the remapping of receptive fields around saccade onset, combining pre- and postsaccadic information. The mislocalization of stimuli briefly flashed near the time of saccades has been taken as evidence for remapping. Yet the relationship between remapping, mislocalization, and trans-saccadic integration remains unclear. We asked participants to identify a target stimulus presented around the time of saccade onset, which was immediately visually masked by a postsaccadic stimulus presented in the same spatial location (backward masking). Presenting two rapidly occurring events across separate fixations allowed us to investigate how the visual system reconstructs what happens during a saccade. We show that saccadic remapping resulted in perception of target and mask as either spatially segregated or integrated, depending on the exact timing of saccade onset. During segregation, the target was unmasked because it was perceived as displaced from the mask; during integration, the postsaccadic stimulus masked the presaccadic target (spatiotopic masking). Thus, segregation and integration may work together to yield continuity in conscious vision.

Contingent adaptation reveals that color is processed faster than motion

Recently is has been suggested that color is processed faster than motion (Zeki & Bartels, Proc. R. Soc. Lond. B 265, 1583 - 1585 . To test this we have made use of color-contingent motion adaptation. The stimulus was a radial luminance-modulated grating. During adaptation the grating oscillated between being green-black and red-black and between rotating clockwise and anti-clockwise. The period of oscillations was 1 sec. The phasic relationship between changes in color and direction of motion was manipulated. At a phasic relationship of 0, clockwise rotation was always green-black and anticlockwise was always red-black. At a phasic relationship of 180 this situation was reversed. A range of phasic relationships from 0 to 324 in steps of 36 (which corresponds to steps of 100ms) was used. At a phasic relationship of 108 degrees, clockwise rotation was physically paired with the red-black color state for 300ms, and with the green-black color state for only 200ms. Yet the contingent aftereffects observed were consistent with clockwise rotation being perceptually correlated more highly with the green-black color state.The reverse situation was observed at a phasic relationship of 288. This indicates that changes in color are processed more than 50ms faster than changes in direction. This is consistent with the hypothesis that the evident processing asynchrony results in an asynchrony of visual consciousness.

Interocular rivalry revealed in the human cortical blind-spot representation

To understand conscious vision, scientists must elucidate how the brain selects specific visual signals for awareness. When different monocular patterns are presented to the two eyes, they rival for conscious expression such that only one monocular image is perceived at a time1,2. Controversy surrounds whether this binocular rivalry reflects neural competition among pattern representations or monocular channels3,4. Here we show that rivalry arises from interocular competition, using functional magnetic resonance imaging of activity in a monocular region of primary visual cortex corresponding to the blind spot. This cortical region greatly prefers stimulation of the ipsilateral eye to that of the blind-spot eye. Subjects reported their dominant percept while viewing rivalrous orthogonal gratings in the visual location corresponding to the blind spot and its surround. As predicted by interocular rivalry, the monocular blind-spot representation was activated when the ipsilateral grating became perceptually dominant and suppressed when the blind-spot grating became dominant. These responses were as large as those observed during actual alternations between the gratings, indicating that rivalry may be fully resolved in monocular visual cortex. Our findings provide the first physiological evidence, to our knowledge, that interocular competition mediates binocular rivalry, and indicate that V1 may be important in the selection and expression of conscious visual information.

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.MethodIn 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.ResultsBlood 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.

Consciousness and attention: on sufficiency and necessity.

Recent research has slowly corroded a belief that selective attention and consciousness are so tightly entangled that they cannot be individually examined. In this review, we summarize psychophysical and neurophysiological evidence for a dissociation between top-down attention and consciousness. The evidence includes recent findings that show subjects can attend to perceptually invisible objects. More contentious is the finding that subjects can become conscious of an isolated object, or the gist of the scene in the near absence of top-down attention; we critically re-examine the possibility of “complete” absence of top-down attention. We also cover the recent flurry of studies that utilized independent manipulation of attention and consciousness. These studies have shown paradoxical effects of attention, including examples where top-down attention and consciousness have opposing effects, leading us to strengthen and revise our previous views. Neuroimaging studies with EEG, MEG, and fMRI are uncovering the distinct neuronal correlates of selective attention and consciousness in dissociative paradigms. These findings point to a functional dissociation: attention as analyzer and consciousness as synthesizer. Separating the effects of selective visual attention from those of visual consciousness is of paramount importance to untangle the neural substrates of consciousness from those for attention.

Does subliminal visual perception have an error‐monitoring system?

There is substantial evidence that subliminal stimuli, i.e. stimuli that cannot be perceived consciously, may influence visually guided human behaviour. Two important points require further investigation, namely, the neural bases and the functional capability of unconscious stimulus processing. In this study we aimed at studying one specific aspect of the latter question, i.e. whether unconscious vision can feed into an error-monitoring system in much the same way as conscious vision. We therefore tested whether the event-related potential component known as error-related negativity, which represents the electrophysiological correlate of an error-monitoring system, is produced by unconscious errors. We found an error-related negativity not only for errors committed in a discrimination task with visible stimuli but also for those committed with subthreshold stimuli. Moreover, behavioural analysis showed post-error slowing of reaction time for correct responses following unconscious as well as conscious errors. Thus, the present results provide both electrophysiological and behavioural evidence of an error-monitoring system operating even when stimuli cannot access consciousness.

How chromatic phenomenality largely overflow its cognitive accessibility

It has been suggested (Block, 2007) that the core neural bases for visual phenomenal consciousness and for access consciousness are located in anatomically separate regions. If this is correct, and if, as Block suggests, the core neural substrate of visual phenomenality is located early in the visual cortex where detailed chromatic information is available, then it would be reasonable to infer that our intuitions of chromatically rich visual phenomenality are plausible. It is furthermore suggested that during perception cognitive access to this chromatic cornucopia is mediated through mereologically superordinate (MS) concepts that regionally characterize both semantic and quantitative integrated properties within complex visual percepts. Such concepts contain much less information than do the particulars that they characterize, implying that the information represented in phenomenal consciousness greatly exceeds the information in the accompanying access consciousness.

“Real-time” obstacle avoidance in the absence of primary visual cortex

When we reach toward objects, we easily avoid potential obstacles located in the workspace. Previous studies suggest that obstacle avoidance relies on mechanisms in the dorsal visual stream in the posterior parietal cortex. One fundamental question that remains unanswered is where the visual inputs to these dorsal-stream mechanisms are coming from. Here, we provide compelling evidence that these mechanisms can operate in "real-time" without direct input from primary visual cortex (V1). In our first experiment, we used a reaching task to demonstrate that an individual with a dense left visual field hemianopia after damage to V1 remained strikingly sensitive to the position of unseen static obstacles placed in his blind field. Importantly, in a second experiment, we showed that his sensitivity to the same obstacles in his blind field was abolished when a short 2-s delay (without vision) was introduced before reach onset. These findings have far-reaching implications, not only for our understanding of the time constraints under which different visual pathways operate, but also in relation to how these seemingly "primitive" subcortical visual pathways can control complex everyday behavior without recourse to conscious vision.

Egocentric Spatial Representation in Action and Perception*

Neuropsychological findings used to motivate the “two visual systems” hypothesis have been taken to endanger a pair of widely accepted claims about spatial representation in conscious visual experience. The first is the claim that visual experience represents 3‐D space around the perceiver using an egocentric frame of reference. The second is the claim that there is a constitutive link between the spatial contents of visual experience and the perceiver’s bodily actions. In this paper, I review and assess three main sources of evidence for the two visual systems hypothesis. I argue that the best interpretation of the evidence is in fact consistent with both claims. I conclude with some brief remarks on the relation between visual consciousness and rational agency.

Temporal constraints on conscious vision: On the ubiquitous nature of the attentional blink

The attentional blink (AB) refers to the finding that observers often miss the second of two masked visual targets (T1 and T2, e.g., letters) appearing within 200-500 ms. Although the presence of a T1 mask is thought to be required for this effect, we recently found that an AB deficit can be observed even in the absence of a T1 mask if T2 is shown very briefly and followed by a pattern mask (M. R. Nieuwenstein, M. C. Potter, & J. Theeuwes, 2009). Using such a sensitive T2 task, the present study sought to determine the minimum requirements for eliciting an AB deficit. To this end, we examined if the occurrence of an AB depends on T1 exposure duration, the requirement to perform a task for T1, and awareness of T1. The results showed that an AB deficit occurs regardless of the presentation duration of T1, and regardless of whether there is a T1 task. A boundary condition for the occurrence of an AB was found in conscious awareness of T1. With a near-threshold detection task for T1, attention blinked when T1 was seen, but not when T1 was missed. Accordingly, we conclude that the minimum requirement for an AB deficit is T1 awareness.