Dissociable rhythmic mechanisms enhance memory for conscious and nonconscious perceptual contents.

Understanding the neural mechanisms of conscious and unconscious experience is a major goal of fundamental and translational neuroscience. Here, we target the early visual cortex with a protocol of noninvasive, high-resolution alternating current stimulation while participants performed a delayed target-probe discrimination task and reveal dissociable mechanisms of mnemonic processing for conscious and unconscious perceptual contents. Entraining ß-rhythms in bilateral visual areas preferentially enhanced short-term memory for seen information, whereas α-entrainment in the same region preferentially enhanced short-term memory for unseen information. The short-term memory improvements were frequency-specific and long-lasting. The results add a mechanistic foundation to existing theories of consciousness, call for revisions to these theories, and contribute to the development of nonpharmacological therapeutics for improving visual cortical processing.

How (not) to demonstrate unconscious priming: Overcoming issues with post-hoc data selection, low power, and frequentist statistics.

One widely used scientific approach to studying consciousness involves contrasting conscious operations with unconscious ones. However, challenges in establishing the absence of conscious awareness have led to debates about the extent and existence of unconscious processes. We collected experimental data on unconscious semantic priming, manipulating prime presentation duration to highlight the critical role of the analysis approach in attributing priming effects to unconscious processing. We demonstrate that common practices like post-hoc data selection, low statistical power, and frequentist statistical testing can erroneously support claims of unconscious priming. Conversely, adopting best practices like direct performance-awareness contrasts, Bayesian tests, and increased statistical power can prevent such erroneous conclusions. Many past experiments, including our own, fail to meet these standards, casting doubt on previous claims about unconscious processing. Implementing these robust practices will enhance our understanding of unconscious processing and shed light on the functions and neural mechanisms of consciousness.

Pupil dilation reflects covert familiar face recognition under interocular suppression.

In prosopagnosia, brain lesions impair overt face recognition, but not face detection, and may coexist with residual covert recognition of familiar faces. Previous studies that simulated covert recognition in healthy individuals have impaired face detection as well as recognition, thus not fully mirroring the deficits in prosopagnosia. We evaluated a model of covert recognition based on continuous flash suppression (CFS). Familiar and unfamiliar faces and houses were masked while participants performed two discrimination tasks. With increased suppression, face/house discrimination remained largely intact, but face familiarity discrimination deteriorated. Covert recognition was present across all masking levels, evinced by higher pupil dilation to familiar than unfamiliar faces. Pupil dilation was uncorrelated with overt performance across subjects. Thus, CFS can impede overt face recognition without disrupting covert recognition and face detection, mirroring critical features of prosopagnosia. CFS could be used to uncover shared neural mechanisms of covert recognition in prosopagnosic patients and neurotypicals.

Using virtual reality to induce multi-trial inattentional blindness despite trial-by-trial measures of awareness.

Unconscious processing has been widely examined using diverse and well-controlled methodologies. However, the extent to which these findings are relevant to real-life instances of information processing without awareness is limited. Here, we present a novel inattentional blindness (IB) paradigm in virtual reality (VR). In three experiments, we managed to repeatedly induce IB while participants foveally viewed salient stimuli for prolonged durations. The effectiveness of this paradigm demonstrates the close relationship between top-down attention and subjective experience. Thus, this method provides an ecologically valid setup to examine processing without awareness.

Pre-stimulus alpha activity modulates long-lasting unconscious feature integration.

Pre-stimulus alpha (α) activity can influence perception of shortly presented, low-contrast stimuli. The underlying mechanisms are often thought to affect perception exactly at the time of presentation. In addition, it is suggested that α cycles determine temporal windows of integration. However, in everyday situations, stimuli are usually presented for periods longer than ∼100 ms and perception is often an integration of information across space and time. Moving objects are just one example. Hence, the question is whether α activity plays a role also in temporal integration, especially when stimuli are integrated over several α cycles. Using electroencephalography (EEG), we investigated the relationship between pre-stimulus brain activity and long-lasting integration in the sequential metacontrast paradigm (SQM), where two opposite vernier offsets, embedded in a stream of lines, are unconsciously integrated into a single percept. We show that increases in α power, even 300 ms before the stimulus, affected the probability of reporting the first offset, shown at the very beginning of the SQM. This effect was mediated by the systematic slowing of the α rhythm that followed the peak in α power. No phase effects were found. Together, our results demonstrate a cascade of neural changes, following spontaneous bursts of α activity and extending beyond a single moment, which influences the sensory representation of visual features for hundreds of milliseconds. Crucially, as feature integration in the SQM occurs before a conscious percept is elicited, this also provides evidence that α activity is linked to mechanisms regulating unconscious processing.

Extracting probability in the absence of visual awareness.

Extracting statistical regularities from the environment is crucial for survival. It allows us to learn cues for where and when future events will occur. Can we learn these associations even when the cues are not consciously perceived? Can these unconscious processes integrate information over long periods of time? We show that human visual system can track the probability of location contingency between an unconscious prime and a conscious target over a period of time of minutes. In a series of psychophysical experiments, we adopted an exogenous priming paradigm and manipulated the location contingency between a masked prime and a visible target (i.e., how likely the prime location predicted the target location). The prime's invisibility was verified both subjectively and objectively. Although the participants were unaware of both the existence of the prime and the prime-target contingency, our results showed that the probability of location contingency was tracked and manifested in the subsequent priming effect. When participants were first entrained into the fully predictive prime-target probability, they exhibited faster responses to the more predictive location. On the contrary, when no contingency existed between the prime and target initially, participants later showed faster responses to the less predictive location. These results were replicated in two more experiments with increased statistical power and a fine-grained delineation of prime awareness. Together, we report that the human visual system is capable of tracking unconscious probability over a period of time, demonstrating how implicit and uncertain regularity guides behavior.

Invisible social grouping facilitates the recognition of individual faces.

Emerging evidence suggests a specialized mechanism supporting perceptual grouping of social entities. However, the stage at which social grouping is processed is unclear. Through four experiments, here we showed that participants' recognition of a visible face was facilitated by the presence of a second facing (thus forming a social grouping) relative to a nonfacing face, even when the second face was invisible. Using a monocular/dichoptic paradigm, we further found that the social grouping facilitation effect occurred when the two faces were presented dichoptically to different eyes rather than monocularly to the same eye, suggesting that social grouping relies on binocular rather than monocular neural channels. The above effects were not found for inverted face dyads, thereby ruling out the contribution of nonsocial factors. Taken together, these findings support the unconscious influence of social grouping on visual perception and suggest an early origin of social grouping processing in the visual pathway.

The role of conscious perception in semantic processing: Testing the action trigger hypothesis.

Finding that invisible primes affect categorization of visible targets (response priming) is held to demonstrate that semantic processing does not require conscious perception. However, the effects are typically very small, they do not indicate whether conscious perception enhances response priming and they often reflect visuo-motor rather than semantic processing. Here, we compared response priming elicited by liminal words when these were clearly seen vs missed, while participants categorized target animals' names. We varied task demands to induce visuo-motor vs semantic processing. Conscious perception strongly enhanced both visuo-motor and semantic response priming. In line with the Action Trigger Hypothesis, task demands modulated processing of both missed and consciously perceived primes. Finally, conscious and unconscious response priming showed diverging patterns on fast and on slow trials, a dissociation suggesting that priming was not contaminated by conscious priming. We conclude that the impact of unconscious stimuli is small and task-dependent.

Neural mechanisms of response priming do not support veridical unconscious processing.

Studies using the priming paradigm often infer that unconscious processes have more veridical access to the world than conscious processes. These interpretations are based on a standard reasoning that erroneously infers good sensitivity of indirect measures from a clear priming effect. To correct for this fallacy, researchers should explicitly compute the sensitivities from indirect measures and compare them against the sensitivities of direct measures. Recent results suggest that indirect behavioral measures are not more sensitive than direct measures and challenge interpretations about veridical unconscious processing. We add to these behavioral results by focusing on neurophysiological indirect measures. In two EEG experiments, we investigate whether event related potentials (ERPs) are more sensitive to different visual stimuli than direct measures. The results show the opposite effect: higher sensitivities for direct than indirect measures. Therefore-contrasting commonly held belief-we find no evidence for more veridical unconscious than conscious processes in ERP measures.

Unconsciously registered items reduce working memory capacity.

The assumption that the contents of consciousness correspond to those of working memory (WM) is challenged by evidence that stimuli masked from awareness can be retained for several seconds (Soto et al., 2011; Bergström & Eriksson, 2015). To assess whether conscious and unconscious items compete in a unitary WM store we conducted an experiment in which some of the memory items in an array were masked from conscious sight using continuous flash suppression (CFS) while others remained visible. After a retention interval, participants decided whether the probed item (either masked or visible) had changed its orientation. Behavioral results indicated that change detection for visible items was significantly impaired when masked items were present, suggesting that masked items either displaced or reduced the precision of visible items in WM. However, change detection for masked items was at chance levels, indicating that these items were not stored. The unsuccessful attempt to encode them may have drawn upon a common pool of attentional resources needed to retain or retrieve visible items. Contralateral Delay Activity, an EEG index of net WM load, failed to temporally localize this interference.

Priming of natural scene categorization during continuous flash suppression.

Continuous Flash Suppression (CFS) reduces conscious awareness of stimuli. Whether stimuli suppressed by CFS are processed at categorical or semantic levels is still debated. Here, we approached this question using a large set of indoor and outdoor scene photographs in a priming paradigm. Perceptually suppressed primes were followed by visible targets. Participants rapidly reported whether the targets showed an indoor or an outdoor scene. Responses were faster (and fast responses more accurate) when primes and targets came from a congruent superordinate category (e.g., both were outdoor scenes). During CFS, priming effects were relatively small (up to 10 ms) and modulated by prime visibility and stimulus onset asynchrony (SOA) of prime and target. Without CFS, the stimuli elicited consistent and more robust priming effects (about 24 ms). Our results imply that scene category is processed during CFS, although some residual prime visibility is likely necessary for significant priming effects to occur.

Recruitment of newly acquired category exemplars into unconscious processing systems is rapid and durable.

In a series of 3 unconscious priming experiments, we investigated if newly acquired English language words can become integrated into unconscious processing systems, and what preconditions are required to enable this process. In each experiment participants learned English language names of extremely rare fish and flowers in a single learning session. After a varying interval of time and in some cases a brief session of repeated conscious exposure (relearning), the newly learned words were presented as briefly flashed, visually masked primes in a standard unconscious category priming procedure. Results show that newly acquired words are recruited quickly into unconscious processing systems. Furthermore, this acquisition persists for at least 48 h and is dependent on prime duration. Since priming was only obtained after an interval that included overnight sleep, consolidation during sleep may play a role in the integration of novel words into unconscious processing systems.

Unconscious semantic priming from pictures under backward masking and continuous flash suppression.

It is debated whether the meaning of invisible pictures can be processed unconsciously. We tested whether pictures of animals or objects presented under backward masking or continuous flash suppression could prime the subsequent categorization of target words into animal or non-animal. In Experiment 1, the backward masking part failed to replicate the priming effect reported in two previous studies, despite sufficient statistical power (N = 59). Similarly, the continuous flash suppression part provided no evidence for a priming effect. In Experiment 2 (N = 65) we shortened the prime-target SOA from 290 ms to 90 ms, but again failed to obtain unconscious semantic priming under backward masking. Thus, our study did not provide evidence for unconscious semantic processing of pictures. These findings support the emerging view that unconscious processing is rather limited in scope.

Back to the future? How Chinese-English bilinguals switch between front and back orientation for time.

The ability to conceive time is a corner stone of human cognition. It is unknown, however, whether time conceptualisation differs depending on language of operation in bilinguals. Whilst both Chinese and English cultures associate the future with the front space, some temporal expressions of Chinese involve a configuration reversal due to historic reasons. For instance, Chinese refers to the day after tomorrow using the spatiotemporal metaphor hou-tian - 'back-day' and to the day before yesterday using qian-tian - 'front-day'. Here, we show that native metaphors interfere with time conceptualisation when bilinguals operate in the second language. We asked Chinese-English bilinguals to indicate whether an auditory stimulus depicted a day of the week either one or two days away from the present day, irrespective of whether it referred to the past or the future, and ignoring whether it was presented through loudspeakers situated in the back or the front space. Stimulus configurations incongruent with spatiotemporal metaphors of Chinese (e.g., "Friday" presented in the front of the participant during a session held on a Wednesday) were conceptually more challenging than congruent configurations (e.g., the same stimulus presented in their back), as indexed by N400 modulations of event-related brain potentials. The same pattern obtained for days or years as stimuli, but surprisingly, it was found only when participants operated in English, not in Chinese. We contend that the task was easier and less prone to induce cross-language activation when conducted in the native language. We thus show that, when they operate in the second language, bilinguals unconsciously retrieve irrelevant native language representations that shape time conceptualisation in real time.

Unconscious Detection of One's Own Image.

A key mechanism behind preferential processing of self-related information might be an early and automatic capture of attention. Therefore, the present study tested a hypothesis that one's own face will attract bottom-up attention even without conscious identification. To test this, we used a dot-probe paradigm with electrophysiological recordings, in which participants ( N = 18) viewed masked and unmasked pairs of faces (other, self) presented laterally. Analysis of the sensitivity measure d ' indicated that faces were not consciously identified in the masked condition. A clear N2 posterior-contralateral (N2pc) component (a neural marker of attention shifts) was found in both the masked and unmasked conditions, revealing that one's own face automatically captures attention when processed unconsciously. Therefore, our study (a) demonstrates that self-related information is boosted at an early (preconscious) stage of processing, (b) identifies further features (beyond simple physical ones) that cause automatic attention capture, and (c) provides further evidence for the dissociative nature of attention and consciousness.

Involuntary mental rotation and visuospatial imagery from external control.

The Reflexive Imagery Task (RIT) was developed to investigate the entry into consciousness of involuntary imagery. Subjects are presented with objects and instructed to not think of the names of the objects. Involuntary subvocalizations arise on many trials. RIT effects reveal the capacities of involuntary processing. These cognitions do not require symbol manipulation. Can mental rotation and visuospatial imagery, too, arise in this involuntary manner? In the mental rotation task, subjects were first taught to mentally rotate two-dimensional objects. Subjects were then instructed to not mentally rotate objects. In the chess task, subjects were taught how to move in their minds objects in specified ways, much as one could imagine how chess pieces move on a chessboard. Subjects were then instructed to not have such visuospatial imagery. For both tasks, involuntary imagery occurred on a substantial proportion of trials, revealing that symbol manipulation can be influenced involuntarily through external control.

Do semantic priming and retrieval of stimulus-response associations depend on conscious perception?

What function does conscious perception serve in human behavior? Many studies relied on unconscious priming to demonstrate that unseen stimuli can be extensively processed. However, showing a small unconscious priming effect falls short of showing that the process underlying such priming is independent of conscious perception. Here, we investigated to what extent the retrieval of learned stimulus-response associations and semantic priming depend on conscious perception by using a liminal-prime paradigm that allows comparing conscious and unconscious processing under the same stimulus conditions. The results revealed two striking dissociations. First, S-R priming was entirely independent of conscious perception, whereas semantic processing was strongly enhanced by it. Second, while priming emerged on fast trials for all conditions, only conscious semantic priming was observed on slow trials. The implications of these findings for the time course of response priming and for the contribution of unconscious processes to fast vs. slow responses are discussed.

"Real-life" continuous flash suppression (CFS)-CFS with real-world objects using augmented reality goggles.

Continuous flash suppression (CFS) is a popular method for suppressing visual stimuli from awareness for relatively long periods. Thus far, this method has only been used for suppressing two-dimensional images presented on screen. We present a novel variant of CFS, termed "real-life" CFS, in which a portion of the actual immediate surroundings of an observer-including three-dimensional, real-life objects-can be rendered unconscious. Our method uses augmented reality goggles to present subjects with CFS masks to the dominant eye, leaving the nondominant eye exposed to the real world. In three experiments we demonstrated that real objects can indeed be suppressed from awareness for several seconds, on average, and that the suppression duration is comparable to that obtained using classic, on-screen CFS. As supplementary information, we further provide an example of experimental code that can be modified for future studies. This technique opens the way to new questions in the study of consciousness and its functions.

Evidence for Implicit-But Not Unconscious-Processing of Object-Scene Relations.

Is consciousness necessary for integration? Findings of seemingly high-level object-scene integration in the absence of awareness have challenged major theories in the field and attracted considerable scientific interest. Lately, one of these findings has been questioned because of a failure to replicate, yet the other finding was still uncontested. Here, we show that this latter finding-slowed-down performance on a visible target following a masked prime scene that includes an incongruent object-is also not reproducible. Using Bayesian statistics, we found evidence against unconscious integration of objects and scenes. Put differently, at the moment, there is no compelling evidence for object-scene congruency processing in the absence of awareness. Intriguingly, however, our results do suggest that consciously experienced yet briefly presented incongruent scenes take longer to process, even when subjects do not explicitly detect their incongruency.

Role of consciousness in temporal integration of semantic information.

Previous studies found that word meaning can be processed unconsciously. Yet it remains unknown whether temporally segregated words can be integrated into a holistic meaningful phrase without consciousness. The first four experiments were designed to examine this by sequentially presenting the first three words of Chinese four-word idioms as prime to one eye and dynamic Mondrians to the other (i.e., the continuous flash suppression paradigm; CFS). An unmasked target word followed the three masked words in a lexical decision task. Results from such invisible (CFS) condition were compared with the visible condition where the preceding words were superimposed on the Mondrians and presented to both eyes. Lower performance in behavioral experiments and larger N400 event-related potentials (ERP) component for incongruent- than congruent-ending words were found in the visible condition. However, no such congruency effect was found in the invisible condition, even with enhanced statistical power and top-down attention, and with several potential confounding factors (contrast-dependent processing, long interval, no conscious training) excluded. Experiment 5 demonstrated that familiarity of word orientation without temporal integration can be processed unconsciously, excluding the possibility of general insensitivity of our paradigm. The overall result pattern therefore suggests that consciousness plays an important role in semantic temporal integration in the conditions we tested.