Working Memory and Consciousness: The Current State of Play.

Working memory (WM), an important posit in cognitive science, allows one to temporarily store and manipulate information in the service of ongoing tasks. WM has been traditionally classified as an explicit memory system-that is, as operating on and maintaining only consciously perceived information. Recently, however, several studies have questioned this assumption, purporting to provide evidence for unconscious WM. In this article, we focus on visual working memory (VWM) and critically examine these studies as well as studies of unconscious perception that seem to provide indirect evidence for unconscious WM. Our analysis indicates that current evidence does not support an unconscious WM store, though we offer independent reasons to think that WM may operate on unconsciously perceived information.

Perceptual overloading reveals illusory contour perception without awareness of the inducers.

Unconscious perception is frequently examined by restricting visual input (e.g., using short stimulus durations followed by masking) to prevent that information from entering visual awareness. Failures to demonstrate perception without awareness may thus be a consequence of this restricted input rather than of limitations in unconscious perception. Here, we demonstrate a novel method that circumvents these significant drawbacks inherent in other methods. Using this new perceptual overloading technique (POT), in which stimuli are repeatedly presented in alternation with a stream of variable masks, we demonstrate illusory contour perception and modal completion even when subjects are completely unaware of the inducing elements. In addition to demonstrating a powerful new method to study consciousness by effectively gating robust visual input from visual awareness, we show that more complex contextual effects, previously considered to be a privilege only of conscious vision, can occur without awareness.

Barack Obama Blindness (BOB): Absence of Visual Awareness to a Single Object.

In two experiments, we evaluated whether a perceiver's prior expectations could alone obliterate his or her awareness of a salient visual stimulus. To establish expectancy, observers first made a demanding visual discrimination on each of three baseline trials. Then, on a fourth, critical trial, a single, salient and highly visible object appeared in full view at the center of the visual field and in the absence of any competing visual input. Surprisingly, fully half of the participants were unaware of the solitary object in front of their eyes. Dramatically, observers were blind even when the only stimulus on display was the face of U.S. President Barack Obama. We term this novel, counterintuitive phenomenon, Barack Obama Blindness (BOB). Employing a method that rules out putative memory effects by probing awareness immediately after presentation of the critical stimulus, we demonstrate that the BOB effect is a true failure of conscious vision.

Feature-based inattentional blindness: loss of awareness to featural information in fully attended objects.

In two experiments, we investigated the impact of feature-based attention on observers' awareness of object appearance. Participants were shown a sequence of two displays, each containing eight objects (rectangles), and were asked to detect changes in the orientation of a cued rectangle. A set of baseline trials preceded probe trials in which half of the rectangles in each display were unexpectedly distorted by 70 %. Participants in both Experiment 1 (100-ms display duration) and Experiment 2 (100- and 400-ms display durations) were unaware of these modifications in the task-irrelevant feature (texture), even when they were asked to select the viewed object in a forced choice procedure. A control experiment showed that participants could identify the physical distortion when they were made aware of its presence. The results demonstrate that feature-based attention moderates the appearance of objects, even when those objects are fully expected and fully attended, implying a distinct form of unawareness that we term feature-based inattentional blindness.

Unconscious priming requires early visual cortex at specific temporal phases of processing.

Although examples of unconscious shape priming have been well documented, whether such priming requires early visual cortex (V1/V2) has not been established. In the current study, we used TMS of V1/V2 at varying temporal intervals to suppress the visibility of preceding shape primes while the interval between primes and targets was kept constant. Our results show that, although conscious perception requires V1/V2, unconscious priming can occur without V1/V2 at an intermediate temporal interval but not at early (5-25 msec) or later (65-125 msec) stages of processing. Because the later time window of unconscious priming suppression has been proposed to interfere with feedback processing, our results further suggest that feedback processing is also essential for unconscious priming and may not be a sufficient condition for conscious vision.

Saliency affects feedforward more than feedback processing in early visual cortex.

Early visual cortex activity is influenced by both bottom-up and top-down factors. To investigate the influences of bottom-up (saliency) and top-down (task) factors on different stages of visual processing, we used transcranial magnetic stimulation (TMS) of areas V1/V2 to induce visual suppression at varying temporal intervals. Subjects were asked to detect and discriminate the color or the orientation of briefly-presented small lines that varied on color saliency based on color contrast with the surround. Regardless of task, color saliency modulated the magnitude of TMS-induced visual suppression, especially at earlier temporal processing intervals that reflect the feedforward stage of visual processing in V1/V2. In a second experiment we found that our color saliency effects were also influenced by an inherent advantage of the color red relative to other hues and that color discrimination difficulty did not affect visual suppression. These results support the notion that early visual processing is stimulus driven and that feedforward and feedback processing encode different types of information about visual scenes. They further suggest that certain hues can be prioritized over others within our visual systems by being more robustly represented during early temporal processing intervals.

Iconic memory requires attention.

Two experiments investigated whether attention plays a role in iconic memory, employing either a change detection paradigm (Experiment 1) or a partial-report paradigm (Experiment 2). In each experiment, attention was taxed during initial display presentation, focusing the manipulation on consolidation of information into iconic memory, prior to transfer into working memory. Observers were able to maintain high levels of performance (accuracy of change detection or categorization) even when concurrently performing an easy visual search task (low load). However, when the concurrent search was made difficult (high load), observers' performance dropped to almost chance levels, while search accuracy held at single-task levels. The effects of attentional load remained the same across paradigms. The results suggest that, without attention, participants consolidate in iconic memory only gross representations of the visual scene, information too impoverished for successful detection of perceptual change or categorization of features.

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.

MecA dampens transitions to spore, biofilm exopolysaccharide and competence expression by two different mechanisms.

The adapter protein MecA targets the transcription factor ComK for degradation by the ClpC/ClpP proteolytic complex, thereby negatively regulating competence in Bacillus subtilis. Here we show that MecA also decreases the frequency of transitions to the sporulation pathway as well as the expression of eps, which encodes synthesis of the biofilm matrix exopolysaccharide. We present genetic and biophysical evidence that MecA downregulates eps expression and spore formation by directly interacting with Spo0A. MecA does not target Spo0A for degradation, and apparently does not prevent the phosphorylation of Spo0A. We propose that it inhibits the transcriptional activity of Spo0A∼P by direct binding. Thus, in its interaction with Spo0A, MecA differs from its role in the regulation of competence where it targets ComK for degradation. MecA acts as a general buffering protein for development, acting by two distinct mechanisms to regulate inappropriate transitions to energy-intensive pathways.

Microarray analysis of the Bacillus subtilis K-state: genome-wide expression changes dependent on ComK.

In Bacillus subtilis, the competence transcription factor ComK activates its own transcription as well as the transcription of genes that encode DNA transport proteins. ComK is expressed in about 10% of the cells in a culture grown to competence. Using DNA microarrays representing approximately 95% of the protein-coding open reading frames in B. subtilis, we compared the expression profiles of wild-type and comK strains, as well as of a mecA mutant (which produces active ComK in all the cells of the population) and a comK mecA double mutant. In these comparisons, we identified at least 165 genes that are upregulated by ComK and relatively few that are downregulated. The use of reporter fusions has confirmed these results for several genes. Many of the ComK-regulated genes are organized in clusters or operons, and 23 of these clusters are preceded by apparent ComK-box promoter motifs. In addition to those required for DNA uptake, other genes that are upregulated in the presence of ComK are probably involved in DNA repair and in the uptake and utilization of nutritional sources. From this and previous work, we conclude that the ComK regulon defines a growth-arrested state, distinct from sporulation, of which competence for genetic transformation is but one notable feature. We suggest that this is a unique adaptation to stress and that it be termed the 'K-state'.

A MecA paralog, YpbH, binds ClpC, affecting both competence and sporulation.

ComK, the master regulator of competence, is degraded by the general stress-related protease ClpCP but must be targeted to this protease by binding to the adapter protein MecA. The genome of Bacillus subtilis contains a paralog of mecA, ypbH. We show in the present study that YpbH, like MecA, binds ClpC and that its elimination or overproduction affects competence and sporulation.