Time, action, and consciousness

Time plays a central role in consciousness, at different levels and in different aspects of information processing. Subliminal perception experiments demonstrate that stimuli presented too briefly to enter conscious awareness are nevertheless processed to some extent. Implicit learning, implicit memory, and conditioning studies suggest that the extent to which memory traces are available for verbal report and for cognitive control is likewise dependent on the time available for processing during acquisition. Differences in the time available for processing also determine not only the extent to which one becomes conscious of action, but also provides the basis for making attributions of authorship to experienced acts. In this paper, we offer a brief overview of these different findings and suggest that they can all be understood based on the fact that consciousness takes time. From this perspective, the availability of representations to conscious awareness depends on the quality of these representations - the extent to which they are strong, stable in time, and distinctive. High-quality representations occur when processes of global competition have had sufficient time to operate so as to make the system settle into the best possible interpretation of the input. Such processes implement global constraint satisfaction and critically depend on reentrant processing, through which representations can be further enriched by high-level constraints. We discuss these ideas in light of current theories of consciousness, emphasizing the fact that consciousness should be viewed as a process rather than as a static property associated with some states and not with others.     

Perceptual awareness and categorical representation of faces: evidence from masked priming

How internal categories influence how we perceive the world is a fundamental question in cognitive sciences. Yet, the relation between perceptual awareness and perceptual categorization has remained largely uncovered so far. Here, we addressed this question by focusing on face perception during subliminal and conscious perception. We used morphed continua between two face identities and we assessed, through a masked priming paradigm, the perceptual processing of these morphed faces under subliminal and supraliminal conditions. We found that priming from subliminal faces followed linearly the information present in the primes, while priming from visible faces revealed a non-linear profile, indicating a categorical processing of face identities. Our results thus point to a special relation between perceptual awareness and categorical processing of faces, and support the dissociation between two modes of information processing: a subliminal mode involving analog treatment of stimuli information, and a supraliminal mode relying on discrete representation.     

Links between conscious awareness and response inhibition: evidence from masked priming

Recent results from “subliminal priming” experiments have shown that masked prime stimuli which can not be consciously perceived can trigger response activation processes, but that these response activations can later be subject to inhibition. Links between conscious awareness and response inhibition were investigated by manipulating the visibility of masked prime stimuli, from clearly visible primes to prime stimuli that were inaccessible to conscious perception. Response inhibition was observed with unperceived prime stimuli, but not for suprathreshold primes. Correlations between individual prime identification thresholds and the onset of response inhibition indicate that the absence or presence of conscious awareness can predict whether or not response inhibition is elicited. These results demonstrate qualitative differences in the effects of conscious and unconscious information. It is argued that response facilitation produced by consciously available perceptual information can counteract automatic effects of self-inhibitory motor control circuits.     

Context-dependent brightness priming occurs without visual awareness

Our visual systems account for stimulus context in brightness perception, but whether such adjustments occur for stimuli that we are unaware of has not been established. We therefore assessed whether stimulus context influences brightness processing by measuring unconscious priming with metacontrast masking. When a middle-gray disk was presented on a darker (or brighter) background, such that it could be consciously perceived as brighter (or darker) via simultaneous brightness contrast (SBC), reaction times were significantly faster to a bright (or dark) annulus than to a dark (or bright) annulus. We further show that context-dependent brightness priming does not correlate with visibility using an objective measure of awareness (Experiment 1) and that context-dependent, but not context-independent brightness priming, occurs equally strongly for stimuli below or above the subjective threshold for awareness (Experiment 2). These results suggest that SBC occurs at early levels of visual input and is not influenced by conscious perception.     

Perception without awareness: perspectives from cognitive psychology

Four basic approaches that have been used to demonstrate perception without awareness are described. Each approach reflects one of two types of experimental logic and one of two possible methods for controlling awareness. The experimental logic has been either to demonstrate a dissociation between a measure of perception with awareness and a measure that is sensitive to perception without awareness or to demonstrate a qualitative difference between the consequences of perception with and without awareness. Awareness has been controlled either by manipulating the stimulus conditions or by instructing observers on how to distribute their attention. The experimental findings based on all four approaches lead to the same conclusion; namely, stimuli are perceived even when observers are unaware of the stimuli. This conclusion is supported by results of studies in which awareness has been assessed with either objective measures of forced-choice discriminations or measures based on verbalizations of subjective conscious experiences. Given this solid empirical support for the concept of perception without awareness, a direction for future research studies is to assess the functions of information perceived without awareness in determining what is perceived with awareness. The available evidence suggests that information perceived without awareness both biases what stimuli are perceived with awareness and influences how stimuli perceived with awareness are consciously experienced.     

Tracking the processes behind conscious perception: a review of event-related potential correlates of visual consciousness

Event-related potential (ERP) studies have attempted to discover the processes that underlie conscious visual perception by contrasting ERPs produced by stimuli that are consciously perceived with those that are not. Variability of the proposed ERP correlates of consciousness is considerable: the earliest proposed ERP correlate of consciousness (P1) coincides with sensory processes and the last one (P3) marks postperceptual processes. A negative difference wave called visual awareness negativity (VAN), typically observed around 200 ms after stimulus onset in occipitotemporal sites, gains strong support for reflecting the processes that correlate with, and possibly enable, aware visual perception. Research suggests that the early parts of conscious processing can proceed independently of top-down attention, although top-down attention may modulate visual processing even before consciousness. Evidence implies that the contents of consciousness are provided by interactions in the ventral stream, but indispensable contributions from dorsal regions influence already low-level visual responses.     

A TMS study of the ventral projections from V1 with implications for the finding of neural correlates of consciousness

The study of subliminal perception in normal and brain lesioned subjects has long been of interest to scholars studying the neural mechanisms behind conscious vision. Using brief durations and a developed methodology of introspective reporting, we present an experiment with visual stimuli that gives rise to little or no subliminal perception under normal viewing conditions. Coupled with transcranial magnetic stimulation, however, we find a dissociation between correctness and conscious awareness. Furthermore, we find support for the hypothesis that the ventral projection streams from V1 are necessary for visual consciousness.     

Working memory biasing of visual perception without awareness

Previous research has demonstrated that the contents of visual working memory can bias visual processing in favor of matching stimuli in the scene. However, the extent to which such top-down, memory-driven biasing of visual perception is contingent on conscious awareness remains unknown. Here we showed that conscious awareness of critical visual cues is dispensable for working memory to bias perceptual selection mechanisms. Using the procedure of continuous flash suppression, we demonstrated that “unseen” visual stimuli during interocular suppression can gain preferential access to awareness if they match the contents of visual working memory. Strikingly, the very same effect occurred even when the visual cue to be held in memory was rendered nonconscious by masking. Control experiments ruled out the alternative accounts of repetition priming and different detection criteria. We conclude that working memory biases of visual perception can operate in the absence of conscious awareness.     

"Binaural rivalry": dichotic listening as a tool for the investigation of the neural correlate of consciousness

Since about two decades neuroscientists have systematically faced the problem of consciousness: the aim is to discover the neural activity specifically related to conscious perceptions, i.e. the biological properties of what philosophers call qualia. In this view, a neural correlate of consciousness (NCC) is a precise pattern of brain activity that specifically accompanies a particular conscious experience. Almost all studies aimed at investigating the NCC have been carried out in the visual system. One of the most promising paradigms is based on sensory stimuli which elicit bistable percepts, as they allow to decouple subjective perception from the characteristics of the physical stimulation. Such kind of perception can be produced in the visual modality by using particular images (e.g. Rubin's vase/face figure) or by presenting two dissimilar stimuli separately to the two eyes (binocular rivalry). The stimuli compete for perceptual dominance and each image is visible in turn for a few seconds, while the other is suppressed. The use of this methodology has led to important findings concerning visual consciousness, which are briefly discussed. For the investigation of auditory consciousness, a similar stimulation paradigm can be achieved by using dichotic listening, consisting in two different stimuli presented each to one ear, which compete for perception (binaural rivalry). The principal aim of the present mini-review is to discuss the few contributes facing the issue of auditory consciousness and to advance the use of dichotic listening and binaural rivalry as valid tools for its investigation.     

Lost in the move? Secondary task performance impairs tactile change detection on the body

Change blindness, the surprising inability of people to detect significant changes between consecutively-presented visual displays, has recently been shown to affect tactile perception as well. Visual change blindness has been observed during saccades and eye blinks, conditions under which people’s awareness of visual information is temporarily suppressed. In the present study, we demonstrate change blindness for suprathreshold tactile stimuli resulting from the execution of a secondary task requiring bodily movement. In Experiment 1, the ability of participants to detect changes between two sequentially-presented vibrotactile patterns delivered on their arms and legs was compared while they performed a secondary task consisting of either the execution of a movement with the right arm toward a visual target or the verbal identification of the target side. The results demonstrated that a motor response gave rise to the largest drop in perceptual sensitivity (as measured by changes in d′) in detecting changes to the tactile display. In Experiment 2, we replicated these results under conditions in which the participants had to detect tactile changes while turning a steering wheel instead. These findings are discussed in terms of the role played by bodily movements, sensory suppression, and higher order information processing in modulating people’s awareness of tactile information across the body surface.     

Prior expectations modulate unconscious evidence accumulation

Unconscious processes have been shown to affect both perception and behaviour. However, the flexibility of such processes remains unknown. Here we investigate whether unconscious decisional processes can adapt to the utility of sensory information. To this end, we had participants gradually accumulate information from noisy motion stimuli, until a decision was reached. We titrated conscious awareness of these stimuli by simultaneously presenting a dynamic dichoptic mask. Crucially, we manipulated the likelihood that the suppressed portion of each presentation would contain useful information. Our results show that the statistics of the environment can be used to modulate unconscious evidence accumulation, resulting in faster choices. Furthermore, computational modelling revealed that this modulation is due to a change in the quality of unconscious evidence accumulation, rather than a conscious change in strategy. Together, these results indicate that unconscious decisional mechanisms are capable of optimising performance by flexibly adapting to the statistical environment.     

Somatosensory processes subserving perception and action

The functions of the somatosensory system are multiple. We use tactile input to localize and experience the various qualities of touch, and proprioceptive information to determine the position of different parts of the body with respect to each other, which provides fundamental information for action. Further, tactile exploration of the characteristics of external objects can result in conscious perceptual experience and stimulus or object recognition. Neuroanatomical studies suggest parallel processing as well as serial processing within the cerebral somatosensory system that reflect these separate functions, with one processing stream terminating in the posterior parietal cortex (PPC), and the other terminating in the insula. We suggest that, analogously to the organisation of the visual system, somatosensory processing for the guidance of action can be dissociated from the processing that leads to perception and memory. In addition, we find a second division between tactile information processing about external targets in service of object recognition and tactile information processing related to the body itself. We suggest the posterior parietal cortex subserves both perception and action, whereas the insula principally subserves perceptual recognition and learning.     

Setting the stage subliminally: unconscious context effects

An important approach to understand how the brain gives rise to consciousness is to probe the depth of unconscious processing, thus to define the key features that cause conscious awareness. Here, we investigate the possibility for subliminal stimuli to shape the context for unconscious processing. Context effects have generally been assumed to require consciousness. In the present experiment, unconscious context processing was investigated by looking at the impact of the context on the response activation elicited by a subliminal prime. We compared the effects of the same subliminal prime on target processing when the prime was embedded in different unconscious contexts. Results showed that the same prime can evoke opposite responses depending on the unconscious context in which it is presented. Taken together, the results of this study show that context effects can be unconscious.     

Empirical support for higher-order theories of conscious awareness

Higher-order theories of consciousness argue that conscious awareness crucially depends on higher-order mental representations that represent oneself as being in particular mental states. These theories have featured prominently in recent debates on conscious awareness. We provide new leverage on these debates by reviewing the empirical evidence in support of the higher-order view. We focus on evidence that distinguishes the higher-order view from its alternatives, such as the first-order, global workspace and recurrent visual processing theories. We defend the higher-order view against several major criticisms, such as prefrontal activity reflects attention but not awareness, and prefrontal lesion does not abolish awareness. Although the higher-order approach originated in philosophical discussions, we show that it is testable and has received substantial empirical support.     

Perception: a model comprising two modes of consciousness

A model of human perception is proposed in which conscious awareness is assumed to be the result of two separate mechanisms each involving consciousness, one outer, sensori-produced, and one inner, conceptual. By mediation of flexible memory representations the sensory data of the outer consciousness give rise to a matched "copy" in the inner consciousness which conceptually organizes the former and also serves as input to the memory store. The model is applied to some perceptual problems in vision such as ambiguous stimuli, subjective contours, space perception, a case of metacontrast, and subliminal perception.     

A hemispheric asymmetry for the unconscious perception of emotion

Previous research has demonstrated that hemispheric asymmetries for conscious visual perception do not lead to asymmetries for unconscious visual perception. These studies utilized emotionally neutral items as stimuli. The current research utilized both emotionally negative and neutral stimuli to assess hemispheric differences for conscious and unconscious visual perception. Conscious perception was measured using a subjective measure of awareness reported by participants on each trial. Unconscious perception was measured by an "exclusion task," a form of word-stem-completion task. Consistent with predictions, negative stimuli were consciously perceived most often when presented to the right hemisphere. Negative stimuli presented to the right hemisphere showed no evidence of unconscious perception, suggesting that the hemispheric asymmetry for the conscious perception of negative information occurs at the expense of unconscious perception.     

To what extent do Gestalt grouping principles influence tactile perception?

Since their formulation by the Gestalt movement more than a century ago, the principles of perceptual grouping have primarily been investigated in the visual modality and, to a lesser extent, in the auditory modality. The present review addresses the question of whether the same grouping principles also affect the perception of tactile stimuli. Although, to date, only a few studies have explicitly investigated the existence of Gestalt grouping principles in the tactile modality, we argue that many more studies have indirectly provided evidence relevant to this topic. Reviewing this body of research, we argue that similar principles to those reported previously in visual and auditory studies also govern the perceptual grouping of tactile stimuli. In particular, we highlight evidence showing that the principles of proximity, similarity, common fate, good continuation, and closure affect tactile perception in both unimodal and crossmodal settings. We also highlight that the grouping of tactile stimuli is often affected by visual and auditory information that happen to be presented simultaneously. Finally, we discuss the theoretical and applied benefits that might pertain to the further study of Gestalt principles operating in both unisensory and multisensory tactile perception.     

Inductive parsimony and the Methodological Argument

Studies on so-called Change Blindness and Inattentional Blindness have been taken to establish the claim that conscious perception of a stimulus requires the attentional processing of that stimulus. One might contend, against this claim, that the evidence only shows attention to be necessary for the subject to have access to the contents of conscious perception and not for conscious perception itself. This "Methodological Argument" is gaining ground among philosophers who work on attention and consciousness, such as Christopher Mole. I find that, without the supporting evidence of inaccessible consciousness, this argument collapses into an indefensible form of inductive parsimony. The Methodological Argument is thus shown to be unsuccessful when used against the claim that attention is required for conscious perception, though I suggest that it may be successful against the more ambitious claim that attention is necessary for all conscious experience.     

Towards a pluralistic neurobiological understanding of consciousness

Theories of consciousness are often based on the assumption that a single, unified neurobiological account will explain different types of conscious awareness. However, recent findings show that, even within a single modality such as conscious visual perception, the anatomical location, timing, and information flow of neural activity related to conscious awareness vary depending on both external and internal factors. This suggests that the search for generic neural correlates of consciousness may not be fruitful. I argue that consciousness science requires a more pluralistic approach and propose a new framework: joint determinant theory (JDT). This theory may be capable of accommodating different brain circuit mechanisms for conscious contents as varied as percepts, wills, memories, emotions, and thoughts, as well as their integrated experience.     

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.