Intracranial neural representation of phenomenal and access consciousness in the human brain.

After more than 30 years of extensive investigation, impressive progress has been made in identifying the neural correlates of consciousness (NCC). However, the functional role of spatiotemporally distinct consciousness-related neural activity in conscious perception is debated. An influential framework proposed that consciousness-related neural activities could be dissociated into two distinct processes: phenomenal and access consciousness. However, though hotly debated, its authenticity has not been examined in a single paradigm with more informative intracranial recordings. In the present study, we employed a visual awareness task and recorded the local field potential (LFP) of patients with electrodes implanted in cortical and subcortical regions. Overall, we found that the latency of visual awareness-related activity exhibited a bimodal distribution, and the recording sites with short and long latencies were largely separated in location, except in the lateral prefrontal cortex (lPFC). The mixture of short and long latencies in the lPFC indicates that it plays a critical role in linking phenomenal and access consciousness. However, the division between the two is not as simple as the central sulcus, as proposed previously. Moreover, in 4 patients with electrodes implanted in the bilateral prefrontal cortex, early awareness-related activity was confined to the contralateral side, while late awareness-related activity appeared on both sides. Finally, Granger causality analysis showed that awareness-related information flowed from the early sites to the late sites. These results provide the first LFP evidence of neural correlates of phenomenal and access consciousness, which sheds light on the spatiotemporal dynamics of NCC in the human brain.

Emotional self-body odors do not influence the access to visual awareness by emotional faces.

A growing body of research suggests that emotional chemosignals in others' body odor (BO), particularly those sampled during fearful states, enhance emotional face perception in conscious and preconscious stages. For instance, emotional faces access visual awareness faster when presented with others' fear BOs. However, the effect of these emotional signals in self-BO, that is, one's own BO, is still neglected in the literature. In the present work, we sought to determine whether emotional self-BOs modify the access to visual awareness of emotional faces. Thirty-eight women underwent a breaking-Continuous Flash Suppression task in which they were asked to detect fearful, happy, and neutral faces, as quickly and accurately as possible, while being exposed to their fear, happiness, and neutral self-BOs. Self-BOs were previously collected and later delivered via an olfactometer, using an event-related design. Results showed a main effect of emotional faces, with happy faces being detected significantly faster than fearful and neutral faces. However, our hypothesis that fear self-BOs would lead to faster emotional face detection was not confirmed, as no effect of emotional self-BOs was found-this was confirmed with Bayesian analysis. Although caution is warranted when interpreting these results, our findings suggest that emotional face perception is not modulated by emotional self-BOs, contrasting with the literature on others' BOs. Further research is needed to understand the role of self-BOs in visual processing and emotion perception.

Mapping the routes of perception: Hemispheric asymmetries in signal propagation dynamics.

The visual system has long been considered equivalent across hemispheres. However, an increasing amount of data shows that functional differences may exist in this regard. We therefore tried to characterize the emergence of visual perception and the spatiotemporal dynamics resulting from the stimulation of visual cortices in order to detect possible interhemispheric asymmetries. Eighteen participants were tested. Each of them received 360 transcranial magnetic stimulation (TMS) pulses at phosphene threshold intensity over left and right early visual areas while electroencephalography was being recorded. After each single pulse, participants had to report the presence or absence of a phosphene. Local mean field power analysis of TMS-evoked potentials showed an effect of both site (left vs. right TMS) of stimulation and hemisphere (ipsilateral vs. contralateral to the TMS): while right TMS determined early stronger activations, left TMS determined later stronger activity in contralateral electrodes. The interhemispheric signal propagation index revealed differences in how TMS-evoked activity spreads: left TMS-induced activity diffused contralaterally more than right stimulation. With regard to phosphenes perception, distinct electrophysiological patterns were found to reflect similar perceptual experiences: left TMS-evoked phosphenes are associated with early occipito-parietal and frontal activity followed by late central activity; right TMS-evoked phosphenes determine only late, fronto-central, and parietal activations. Our results show that left and right occipital TMS elicits differential electrophysiological patterns in the brain, both per se and as a function of phosphene perception. These distinct activation patterns may suggest a different role of the two hemispheres in processing visual information and giving rise to perception.

Attentional blur and blink: Effects of adaptive attentional scaling on visual awareness.

Attentional scaling is a crucial mechanism that enables us to flexibly allocate our attention to larger or smaller regions in the visual field. Although previous studies have demonstrated the critical role of attentional scaling in visual processing, its impact on modulating visual awareness is not yet fully understood. This study investigates the adaptive control of attentional scaling and its influence on visual awareness in an attentional blink paradigm. Participants were required to attend to the first target's location, which was manipulated either session-wise, trial-wise, or such that it could be learned across a block of trials. Discrete, all-or-none, awareness was expected when attention was allocated to a narrow area, while gradual awareness was expected when attention was allocated to a larger area. We used mixture modeling to assess second target awareness across these different attentional scales. The results revealed that participants could adaptively control their attentional scale both across stable sessions, and through (implicit) statistical learning in blocks of successive trials. This produced gradual perceptual awareness when the participants adopted a broad attentional scale, causing an attentional "blur". However, trial-wise cues did not allow for attentional scaling, resulting in more discrete target perception overall, and an attentional "blink". We conclude that the attentional scale is to some extent under adaptive control during the attentional blink/blur, where it can produce qualitatively different modes of perceptual awareness.

Distrust before first sight? Examining knowledge- and appearance-based effects of trustworthiness on the visual consciousness of faces.

The present EEG study with 32 healthy participants investigated whether affective knowledge about a person influences the visual awareness of their face, additionally considering the impact of facial appearance. Faces differing in perceived trustworthiness based on appearance were associated with negative or neutral social information and shown as target stimuli in an attentional blink task. As expected, participants showed enhanced awareness of faces associated with negative compared to neutral social information. On the neurophysiological level, this effect was connected to differences in the time range of the early posterior negativity (EPN)-a component associated with enhanced attention and facilitated processing of emotional stimuli. The findings indicate that the social-affective relevance of a face based on emotional knowledge is accessed during a phase of attentional enhancement for conscious perception and can affect prioritization for awareness. In contrast, no clear evidence for influences of facial trustworthiness during the attentional blink was found.

Dissociation between temporal attention and Consciousness: Unconscious temporal cue induces temporal expectation effect.

The debate over the independence of attention and consciousness is ongoing. Prior studies have established that invisible spatial cues can direct attention. However, our exploration extends beyond spatial dimensions to temporal information as a potent guide for attention. A intriguing question arises: Can unconscious temporal cues trigger attentional orienting? To investigate, we employed a modified reaction-time task in Experiments 1 and 2, using Gabor stimuli or human facial stimuli as temporal cues rendered invisible through continuous flash suppression. We aimed to uncover temporal expectation effects (TE effects) without conscious awareness. Moreover, Experiments 3 and 4 probed the boundaries of this unconscious processing, assessing whether conscious temporal cues could modulate TE effects. Our results confirm that invisible temporal cues can induce TE effects, and these effects can be overridden by conscious temporal cues. This dissociation between temporal attention and consciousness provide a new perspective on our understanding of their relationship.

Prior conscious experience modulates the impact of audiovisual temporal correspondence on unconscious visual processing.

Conscious visual experiences are enriched by concurrent auditory information, implying audiovisual interactions. In the present study, we investigated how prior conscious experience of auditory and visual information influences the subsequent audiovisual temporal integration under the surface of awareness. We used continuous flash suppression (CFS) to render perceptually invisible a ball-shaped object constantly moving and bouncing inside a square frame window. To examine whether audiovisual temporal correspondence facilitates the ball stimulus to enter awareness, the visual motion was accompanied by click sounds temporally congruent or incongruent with the bounces of the ball. In Experiment 1, where no prior experience of the audiovisual events was given, we found no significant impact of audiovisual correspondence on visual detection time. However, when the temporally congruent or incongruent bounce-sound relations were consciously experienced prior to CFS in Experiment 2, congruent sounds yielded faster detection time compared to incongruent sounds during CFS. In addition, in Experiment 3, explicit processing of the incongruent bounce-sound relation prior to CFS slowed down detection time when the ball bounces became later congruent with sounds during CFS. These findings suggest that audiovisual temporal integration may take place outside of visual awareness though its potency is modulated by previous conscious experiences of the audiovisual events. The results are discussed in light of the framework of multisensory causal inference.

Disentangling perceptual awareness from nonconscious processing in rhesus monkeys (Macaca mulatta).

Scholars have long debated whether animals, which display impressive intelligent behaviors, are consciously aware or not. Yet, because many complex human behaviors and high-level functions can be performed without conscious awareness, it was long considered impossible to untangle whether animals are aware or just conditionally or nonconsciously behaving. Here, we developed an empirical approach to address this question. We harnessed a well-established cross-over double dissociation between nonconscious and conscious processing, in which people perform in completely opposite ways when they are aware of stimuli versus when they are not. To date, no one has explored if similar performance dissociations exist in a nonhuman species. In a series of seven experiments, we first established these signatures in humans using both known and newly developed nonverbal double-dissociation tasks and then identified similar signatures in rhesus monkeys (Macaca mulatta). These results provide robust evidence for two distinct modes of processing in nonhuman primates. This empirical approach makes it feasible to disentangle conscious visual awareness from nonconscious processing in nonhuman species; hence, it can be used to strip away ambiguity when exploring the processes governing intelligent behavior across the animal kingdom. Taken together, these results strongly support the existence of both nonconscious processing as well as functional human-like visual awareness in nonhuman animals.

When mind and body align: examining the role of cross-modal congruency in conscious representations of happy facial expressions.

This study explored how congruency between facial mimicry and observed expressions affects the stability of conscious facial expression representations. Focusing on the congruency effect between proprioceptive/sensorimotor signals and visual stimuli for happy expressions, participants underwent a binocular rivalry task displaying neutral and happy faces. Mimicry was either facilitated with a chopstick or left unrestricted. Key metrics included Initial Percept (bias indicator), Onset Resolution Time (time from onset to Initial Percept), and Cumulative Time (content stabilization measure). Results indicated that mimicry manipulation significantly impacted Cumulative Time for happy faces, highlighting the importance of congruent mimicry in stabilizing conscious awareness of facial expressions. This supports embodied cognition models, showing the integration of proprioceptive information significantly biases conscious visual perception of facial expressions.

Continuous Theta Burst Stimulation to the left anterior medial prefrontal cortex influences metacognitive efficiency.

The contribution of the prefrontal areas to visual awareness is critical for the Global Neuronal Workspace Theory and higher-order theories of consciousness. The goal of the present study was to test the potential engagement of the anterior medial prefrontal cortex (aMPFC) in visual awareness judgements. We aimed to temporarily influence the neuronal dynamics of the left aMPFC via neuroplasticity-like mechanisms. We used different Theta Burst Stimulation (TBS) protocols in combination with a visual identification task and visual awareness ratings. Either continuous TBS (cTBS), intermittent TBS (iTBS), or sham TBS was applied prior to the experimental paradigm in a within-participant design. Compared with sham TBS, we observed an increase in participants' ability to judge their perception adequately (metacognitive efficiency) following cTBS but not iTBS. The effect was accompanied by lower visual awareness ratings in incorrect responses. No significant differences in the identification task performance were observed. We interpret these results as evidence of the involvement of PFC in the brain network that underlies metacognition. Further, we discuss whether the results of TMS studies on perceptual metacognition can be taken as evidence for PFC involvement in awareness itself.

Revealing robust neural correlates of conscious and unconscious visual processing: Activation likelihood estimation meta-analyses.

Our ability to consciously perceive information from the visual scene relies on a myriad of intrinsic neural mechanisms. Functional neuroimaging studies have sought to identify the neural correlates of conscious visual processing and to further dissociate from those pertaining to preconscious and unconscious visual processing. However, delineating what core brain regions are involved in eliciting a conscious percept remains a challenge, particularly regarding the role of prefrontal-parietal regions. We performed a systematic search of the literature that yielded a total of 54 functional neuroimaging studies. We conducted two quantitative meta-analyses using activation likelihood estimation to identify reliable patterns of activation engaged by i. conscious (n = 45 studies, comprising 704 participants) and ii. unconscious (n = 16 studies, comprising 262 participants) visual processing during various task performances. Results of the meta-analysis specific to conscious percepts quantitatively revealed reliable activations across a constellation of regions comprising the bilateral inferior frontal junction, intraparietal sulcus, dorsal anterior cingulate, angular gyrus, temporo-occipital cortex and anterior insula. Neurosynth reverse inference revealed conscious visual processing to be intertwined with cognitive terms related to attention, cognitive control and working memory. Results of the meta-analysis on unconscious percepts revealed consistent activations in the lateral occipital complex, intraparietal sulcus and precuneus. These findings highlight the notion that conscious visual processing readily engages higher-level regions including the inferior frontal junction and unconscious processing reliably recruits posterior regions, mainly the lateral occipital complex.

The spectral dynamics of visual awareness: An interplay of different frequencies?

The goal of this study was to investigate the neurophysiological correlates of visual awareness, with a specific focus on its event-related spectral perturbation (ERSP) features. To this aim, we tried to disentangle the proper neural correlates of consciousness (NCC) from other prerequisite and post-perceptual processing. To do so, we administered an orientation discrimination task, inducing a response bias through task instructions. EEG results showed that different frequency bands are involved in this kind of task, with different spectral and temporal dynamics. In particular, alpha and beta bands seem to be particularly engaged, especially in the aware-unaware contrast, showing a main power suppression for aware trials and replicating previous literature. Moreover, we demonstrated that the process of visual awareness is orchestrated by a complex interaction of different frequencies (i.e., theta, alpha, beta and gamma) being involved as prerequisites and post-perceptual processes.

Amplitude modulating frequency overrides carrier frequency in tACS-induced phosphene percept.

The amplitude modulated (AM) neural oscillation is an essential feature of neural dynamics to coordinate distant brain areas. The AM transcranial alternating current stimulation (tACS) has recently been adopted to examine various cognitive functions, but its neural mechanism remains unclear. The current study utilized the phosphene phenomenon to investigate whether, in an AM-tACS, the AM frequency could modulate or even override the carrier frequency in phosphene percept. We measured the phosphene threshold and the perceived flash rate/pattern from 12 human subjects (four females, aged from 20-44 years old) under tACS that paired carrier waves (10, 14, 18, 22 Hz) with different envelope conditions (0, 2, 4 Hz) over the mid-occipital and left facial areas. We also examined the phosphene source by adopting a high-density stimulation montage. Our results revealed that (1) phosphene threshold was higher for AM-tACS than sinusoidal tACS and demonstrated different carrier frequency functions in two stimulation montages. (2) AM-tACS slowed down the phosphene flashing and abolished the relation between the carrier frequency and flash percept in sinusoidal tACS. This effect was independent of the intensity change of the stimulation. (3) Left facial stimulation elicited phosphene in the upper-left visual field, while occipital stimulation elicited equally distributed phosphene. (4) The near-eye electrodermal activity (EDA) measured under the threshold-level occipital tACS was greater than the lowest power sufficient to elicit retinal phosphene. Our results show that AM frequency may override the carrier frequency and determine the perceived flashing frequency of AM-tACS-induced phosphene.

The linear impact of visual working memory load on visual awareness: Evidence from motion-induced blindness.

Despite the close relationship between visual working memory (VWM) and visual awareness, the question of how these two constructs interact with each other is still under debate. The current study aimed to further address this issue by investigating whether and how visual awareness is influenced by VWM load. In Experiment 1, participants were asked to perform a motion-induced blindness (MIB) task while simultaneously memorizing different numbers of items in VWM. The results indicated that the latency of MIB was prolonged gradually as the VWM load increased, revealing a linear trend in the modulation effect of VWM load on visual awareness. Experiments 2 and 3 tested the other potential explanations and validated the initial finding by confirming that VWM load was indeed responsible for the observed effect on visual awareness. These findings have important implications for a better understanding of the relationship between VWM and visual awareness.

Neural correlates of consciousness in an attentional blink paradigm with uncertain target relevance.

Several event-related potentials (ERPs) have been proposed as neural correlates of consciousness (NCC), most prominently the early visual awareness negativity (VAN) and the late P3b component. Highly influential support for the P3b comes from studies utilizing the attentional blink (AB), where conscious perception of a first visual target (T1) impairs reporting a second target (T2) presented shortly afterwards. Recent no-report studies using other paradigms suggest that the P3b component may reflect post-perceptual processes associated with decision-making rather than awareness. However, no-report studies are limited in their awareness assessment, and their conclusions have not been tested in an AB paradigm. The present study (N = 38) addressed these issues using a novel AB paradigm, which reduced decision-making processes by omitting a discrimination task on T2 stimuli and rendering their relevance uncertain. Nevertheless, awareness was assessed trial by trial. Comparing ERPs in response to seen versus unseen T2 stimuli revealed a VAN but no enhanced P3b regardless of whether they were marked as distinct from distractor stimuli or not. Our results corroborate the VAN and challenge the P3b as NCC despite rigorous trial-by-trial assessment of conscious perception. Thus, they support the idea that awareness emerges during early sensory processing.

Conflict monitoring and attentional adjustment during binocular rivalry.

To make sense of ambiguous and, at times, fragmentary sensory input, the brain must rely on a process of active interpretation. At any given moment, only one of several possible perceptual representations prevails in our conscious experience. Our hypothesis is that the competition between alternative representations induces a pattern of neural activation resembling cognitive conflict, eventually leading to fluctuations between different perceptual outcomes in the case of steep competition. To test this hypothesis, we probed changes in perceptual awareness between competing images using binocular rivalry. We drew our predictions from the conflict monitoring theory, which holds that cognitive control is invoked by the detection of conflict during information processing. Our results show that fronto-medial theta oscillations (5-7 Hz), an established electroencephalography (EEG) marker of conflict, increases right before perceptual alternations and decreases thereafter, suggesting that conflict monitoring occurs during perceptual competition. Furthermore, to investigate conflict resolution via attentional engagement, we looked for a neural marker of perceptual switches as by parieto-occipital alpha oscillations (8-12 Hz). The power of parieto-occipital alpha displayed an inverse pattern to that of fronto-medial theta, reflecting periods of high interocular inhibition during stable perception, and low inhibition around moments of perceptual change. Our findings aim to elucidate the relationship between conflict monitoring mechanisms and perceptual awareness.

Perceptual filling-in dispels the veridicality problem of conscious perception research.

Conscious perceptual experiences are expected to correlate with content-specific brain activity. A veridicality problem arises when attempting to disentangle unconscious and conscious brain processes if conscious perceptual contents accurately match the physical nature of the stimulus. We argue that perceptual filling-in, a phenomenon whereby visual information inaccurately spreads across visual space, is a promising approach to circumvent the veridicality problem. Filling-in generates non-veridical although unambiguous percepts dissociated from stimulus input. In particular, the radial uniformity illusion induces a filling-in experience between a central disk and the surrounding periphery. We discuss how this illusion facilitates both the detection of neurophysiological responses and subjective phenomenological monitoring. We report behavioral effects from a large-sample (n = 200) psychophysics study and examine key stimulus parameters that drive the conscious filling-in experience. We propose that these data underpin future hypothesis-driven studies of filling-in to further delineate the neural mechanisms of conscious perception.

Typicality modulates the visual awareness of objects.

In the present study, we assessed whether typicality can influence the visual awareness of objects. Participants tracked moving images of objects and counted how often members of one category bounced off the edges of the display. On the last trial, an unexpected object moved across the display. In our first two experiments, this object could belong to the same category as the tracked or untracked objects. While participants were more likely to notice atypical members of the untracked category, this pattern of results reversed when participants tracked atypical objects. In our last two experiments, the unexpected object could belong to the same category as the tracked objects or a new category of objects. In this case, participants were more likely to notice typical members of both the tracked category and the new category. Together, these findings suggest that typicality can modulate the visual awareness of objects.

Differential Effects of Awareness and Task Relevance on Early and Late ERPs in a No-Report Visual Oddball Paradigm.

To date it is poorly understood how and when deviance processing interacts with awareness and task relevance. Furthermore, an important issue in the study of consciousness is the prevalent confound of conscious perception with the requirement of reporting it. This study addresses these topics using a no-report inattentional blindness paradigm with a visual oddball sequence of geometrical shapes presented to male and female human participants. Electrophysiological responses were obtained in three physically identical Phases A-C that differed only with respect to the instructions: (A) participants were uninformed about the shapes and attended an unrelated foreground task (inattentional blind), (B) were informed about the shapes but still attended the foreground task, and (C) attended the shapes. Conscious processing of shapes was indexed by the visual awareness negativity but not a P3. Deviance processing was associated with the visual mismatch negativity independently of consciousness and task relevance. The oddball P3, however, only emerged when the stimuli were task relevant, and was absent for consciously perceived but task irrelevant stimuli. The P3 thus does not represent a reliable marker of stimulus awareness. This result pattern supports the view of hierarchical predictive processing, where lower levels display automatic deviance processing, whereas higher levels require attention and task relevance.SIGNIFICANCE STATEMENT To react to potentially important changes in our environment it is fundamental to detect deviations from regularities of sensory input. It has yet to be understood how awareness and task relevance of this input interact with deviance processing. We investigated the role of awareness in deviance detection while at the same time circumventing the confound of awareness and report by means of a no-report paradigm. Our results suggest that early processes are elicited automatically, whereas, contrary to prominent theories, late processes do not depend on awareness but on task-based attention.

Conscious awareness modulates processing speed in the redundant signal effect.

Evidence for the influence of unaware signals on behaviour has been reported in both patient groups and healthy observers using the Redundant Signal Effect (RSE). The RSE refers to faster manual reaction times to the onset of multiple simultaneously presented target than those to a single stimulus. These findings are robust and apply to unimodal and multi-modal sensory inputs. A number of studies on neurologically impaired cases have demonstrated that RSE can be found even in the absence of conscious experience of the redundant signals. Here, we investigated behavioural changes associated with awareness in healthy observers by using Continuous Flash Suppression to render observers unaware of redundant targets. Across three experiments, we found an association between reaction times to the onset of a consciously perceived target and the reported level of visual awareness of the redundant target, with higher awareness being associated with faster reaction times. However, in the absence of any awareness of the redundant target, we found no evidence for speeded reaction times and even weak evidence for an inhibitory effect (slowing down of reaction times) on response to the seen target. These findings reveal marked differences between healthy observers and blindsight patients in how aware and unaware information from different locations is integrated in the RSE.