The influence of spatial and temporal attention on visual awareness-a behavioral and ERP study.

Whether attention is a prerequisite of perceptual awareness or an independent and dissociable process remains a matter of debate. Importantly, understanding the relation between attention and awareness is probably not possible without taking into account the fact that both are heterogeneous and multifaceted mechanisms. Therefore, the present study tested the impact on visual awareness of two attentional mechanisms proposed by the Posner model: temporal alerting and spatio-temporal orienting. Specifically, we evaluated the effects of attention on the perceptual level, by measuring objective and subjective awareness of a threshold-level stimulus; and on the neural level, by investigating how attention affects two postulated event-related potential correlates of awareness. We found that alerting and orienting mechanisms additively facilitate perceptual consciousness, with activation of the latter resulting in the most vivid awareness. Furthermore, we found that late positivity is unlikely to constitute a neural correlate of consciousness as its amplitude was modulated by both attentional mechanisms, but early visual awareness negativity was independent of the alerting and orienting mechanisms. In conclusion, our study reveals a nuanced relationship between attention and awareness; moreover, by investigating the effect of the alerting mechanism, this study provides insights into the role of temporal attention in perceptual consciousness.

Early Electrophysiological Correlates of Perceptual Consciousness Are Affected by Both Exogenous and Endogenous Attention.

It has been proposed that visual awareness negativity (VAN), which is an early ERP component, constitutes a neural correlate of visual consciousness that is independent of perceptual and cognitive mechanisms. In the present study, we investigated whether VAN is indeed a specific marker of phenomenal awareness or rather reflects the involvement of attention. To this end, we reanalyzed data collected in a previously published EEG experiment in which awareness of visual stimuli and two aspects that define attentional involvement, namely, the inherent saliency and task relevance of a stimulus, were manipulated orthogonally. During the experimental procedure, participants (n = 41) were presented with images of faces that were backward-masked or unmasked, fearful or neutral, and defined as task-relevant targets or task-irrelevant distractors. Single-trial ERP analysis revealed that VAN was highly dependent on attentional manipulations in the early time window (140-200 msec), up to the point that the effect of awareness was not observed for attentionally irrelevant stimuli (i.e., neutral faces presented as distractors). In the late time window (200-350 msec), VAN was present in all attentional conditions, but its amplitude was significantly higher in response to fearful faces and task-relevant face images than in response to neutral ones and task-irrelevant ones, respectively. In conclusion, we demonstrate that the amplitude of VAN is highly dependent on both exogenous (stimulus saliency) and endogenous attention (task requirements). Our results challenge the view that VAN constitutes an attention-independent correlate of phenomenal awareness.

Naturalistic use of psychedelics does not modulate processing of self-related stimuli (but it might modulate attentional mechanisms): An event-related potentials study.

Classic psychedelics are able to profoundly alter the state of consciousness and lead to acute experiences of ego dissolution - the blurring of the distinction between representations of self and the external world. However, whether repeated use of psychedelics is associated with more prolonged and permanent modifications to the concept of self remains to be investigated. Therefore, we conducted a preregistered, cross-sectional study in which experienced psychedelics users (15 or more lifetime experiences with psychedelics; N = 56) were compared to nonusers (N = 57) in terms of neural reactivity to a Self-name (i.e., each participant's own name) stimulus, which is known to robustly activate a representation of self. Two control stimuli were additionally used: an Other-name stimulus, as a passive control condition in which no reaction was required, and a Target-name stimulus, to which participants provided a manual response and which thus constituted an active control condition. Analysis of the amplitude of the P300 ERP component evoked by the Self- or Target-names revealed no difference between the psychedelics users and nonusers. However, psychedelic users exhibited increased P300 amplitude during perception of Other-names. In addition, in comparison to nonusers, psychedelics users exhibited a smaller increase in P300 amplitude when processing the task-relevant Target-names (in relation to both Self- and Other-names). Therefore, our data suggests that regular naturalistic use of psychedelics may not be related to long-term changes in the representation of self, but it might potentially affect the allocation of attentional resources to task-relevant stimuli.

The relation between naturalistic use of psychedelics and perception of emotional stimuli: An event-related potential study comparing non-users and experienced users of classic psychedelics.

BACKGROUND: Previous research has suggested that controlled administration of psychedelic substances can modulate emotional reactivity, enhancing positive and diminishing negative emotions. However, it is unclear whether similar effects are associated with using psychedelics in less-controlled naturalistic environments. AIMS: This cross-sectional study investigated the neural markers associated with the perception of emotional stimuli in individuals with extensive experience of naturalistic psychedelic use (15 or more lifetime experiences), comparing them to non-users. METHODS: Electroencephalography (EEG) signals were recorded from two groups: experienced psychedelics users (N = 56) and non-users (N = 55). Participants were presented with facial images depicting neutral or emotional expressions (anger, sadness, and happiness). Event-related potential (ERP) components were analyzed as indices of emotional reactivity. RESULTS: Psychedelic users were characterized by significantly lower amplitudes of the N200 component in response to fearful faces, in comparison to non-users. In addition, interaction effects between Group and Emotional expression were observed on N170 and N200 amplitudes, indicating group differences in the processing of fearful faces. However, no significant between-group differences emerged in the analysis of later ERP components associated with attention and cognitive processes (P200 and P300). CONCLUSIONS: The results suggest that naturalistic use of psychedelics may be linked to reduced reactivity to emotionally negative stimuli at the early and automatic processing stages. Our study contributes to a better understanding of the effects related to using psychedelics in naturalistic contexts.

I focus only when I see your fear-fearful faces are not prioritized by attention when processed outside of awareness.

The defensive reaction to threats consists of two components: non-specific physiological arousal and specific attentional prioritization of the threatening stimulus, both of which are assumed by the so-called "low-road" hypothesis to be induced automatically and unconsciously. Although ample evidence indicates that non-specific arousal can indeed be caused by unconscious threatening stimuli, data regarding the involvement of the attentional selection mechanism remain inconclusive. Therefore, in the present study we used ERPs to compare the potential engagement of attention in the perception of subliminal and supraliminal fearful facial expressions to that of neutral ones. In the conscious condition, fearful faces were preferentially encoded (as indicated by the N170 component) and prioritized by bottom-up (EPN) and spatial attention (N2pc) in an automatic, task-independent manner. Furthermore, consciously perceived fearful expressions engaged cognitive resources (SPCN, P3) when face stimuli were task-relevant. In the unconscious condition, fearful faces were still preferentially encoded (N170), but we found no evidence for any type of attentional prioritization. Therefore, by showing that threatening stimuli engage attention only when perceived consciously, our findings challenge the "low road" hypothesis and point to the limits of unconscious attentional selection.

Sensory modality defines the relation between EEG Lempel-Ziv diversity and meaningfulness of a stimulus.

Diversity of brain activity is a robust neural correlate of global states of consciousness. It has been proposed that diversity measures specifically reflect the temporal variability of conscious experience. Previous studies supported this hypothesis by showing that perception of meaningful visual stimuli causes richer, more-variable experiences than perception of meaningless stimuli, and this is reflected in greater brain signal diversity. To investigate whether this relation is consistent across sensory modalities, to participants we presented three versions of naturalistic visual and auditory stimuli (videos and audiobooks) that varied in the amount of meaning (original, scrambled, and noise), while recording electroencephalographic signals. We report three main findings. First, greater meaningfulness of visual stimuli was related to higher Lempel-Ziv diversity of EEG signals, but the opposite effect was found in the auditory modality. Second, visual perception was related to generally higher EEG diversity than auditory perception. Third, perception of meaningful visual stimuli and auditory stimuli respectively resulted in higher and lower EEG diversity in comparison to the resting state. In conclusion, the signal diversity of continuous brain signals depends on the stimulated sensory modality, therefore it is not a generic index of the variability of conscious experience.

Attentional Prioritization of Complex, Naturalistic Stimuli Maintained in Working-Memory-A Dot-Probe Event-Related Potentials Study.

Recent studies suggest that a stimulus actively maintained in working memory (WM) automatically captures visual attention when subsequently perceived. Such a WM-based guidance effect has been consistently observed for stimuli defined by simple features, such as color or orientation, but studies using more complex stimuli provided inconclusive results. Therefore, we investigated whether the WM-based guidance effect occurs also for naturalistic stimuli, whose identity is defined by multiple features and relations among them, specifically for faces and houses. The experiment consisted of multiple blocks in which participants (N = 28) either memorized or merely saw (WM or exposure condition, respectively) a template stimulus and then performed several dot-probe trials, with pairs of stimuli (template and control) presented laterally as distractors and followed by a target-asterisk. Analysis of reaction-times (RT) in the dot-probe task shows that memorized stimuli were prioritized by attention and points toward attention-hold, rather than capture, as a mechanism of attentional prioritization. Consistent with this interpretation, memorized items did not evoke a lateralized N2pc ERP component, thought to indicate attention shifts. However, in an exploratory ERP analysis we found evidence for a very early (100-200 ms post-stimulus) prioritization specific to the memorized faces, which is in line with the sensory recruitment theory of WM. In conclusion, our data provide evidence that complex stimuli are prioritized by attention when maintained in WM, and that the mechanism of such prioritization is based on a prolonged hold of spatial attention.

Naturalistic use of psychedelics is related to emotional reactivity and self-consciousness: The mediating role of ego-dissolution and mystical experiences.

BACKGROUND: Psychedelics are able to acutely alter emotional reactivity and self-consciousness. However, whether the regular naturalistic use of psychedelics can be linked to more persistent trait-level changes in these domains remains an open question. AIM: To test the hypotheses that (1) using psychedelics is related to higher positive and lower negative emotional reactivity; and (2) an adaptive pattern of self-consciousness, including diminished public self-consciousness and rumination, and increased reflection and self-awareness; and (3) these relations are mediated by the intensity of past ego-dissolution and mystical experiences. METHOD: An online survey including questions about the history of psychoactive substance use; questionnaires measuring trait levels of emotional reactivity and self-consciousness; questionnaires for retrospective assessment of ego-dissolution and mystical experiences. Data collected from 2516 participants (1661 psychedelics users) were analyzed using robust linear regression and mediation analysis. RESULTS: A higher number of lifetime uses of psychedelics predicted greater positive and lower negative emotional reactivity; also, in the domain of self-consciousness, it predicted greater reflection and internal state awareness, and reduced rumination tendency and public self-consciousness. Finally, the intensity of past mystical and ego-dissolution experiences mediated almost all the observed relationships between the lifetime number of psychedelics uses and psychological variables. CONCLUSIONS: Lifetime psychedelics use predicts an adaptive pattern of trait-level emotional reactivity and self-consciousness. Ego-dissolution and mystical experiences are essential in understanding the long-lasting psychological effects of psychedelics use. Our findings might potentially explain previous observations of increased well-being in psychedelics users.

Perception of semantic relations in scenes: A registered report study of attention hold.

To what extent the semantic relations present in scenes guide spatial attention automatically remains a matter of debate. Considering that spatial attention can be understood as a sequence of shifts, engagements, and disengagements, semantic relations might affect each stage of this process differently. Therefore, we investigated whether objects that violate semantic rules engage attention for longer than objects that are expected in a given context. The experiment involved a central presentation of a distractor scene that contained a semantically congruent or incongruent object, and a peripheral presentation of a small target letter. We found that incongruent scenes did not delay responses to the peripheral target, which indicates that they did not hold attention for longer than congruent scenes. Therefore, by showing that violations of semantic relations do not engage attention automatically, our study contributes to a better understanding of how attention operates in naturalistic settings.

The forest, the trees, or both? Hierarchy and interactions between gist and object processing during perception of real-world scenes.

The global-to-local theories of perception assume that the gist of a scene is computed early and automatically, whereas recognition of objects occurs at a later processing stage, requires attentional resources, and is primed by the representation of gist. To test these theoretical predictions, we investigated the processing hierarchy of gist- and object-recognition and their interaction in two experiments (total N = 60). Backward-masked images of real-world scenes were presented for a range of brief durations - between 8 ms and 100 ms, and participants performed either an object or a background classification task, in separate blocks. We report three main findings. First, scenes' backgrounds were generally classified more accurately than foreground objects, but recognition of objects was boosted to the same level as backgrounds by cueing spatial attention to the exact object's location. Second, backgrounds influence objects' recognition, as objects presented within semantically incongruent backgrounds were classified less accurately. Third, objects influence background categorization, as backgrounds comprising incongruent objects were also classified less accurately. Therefore, the first two findings support the global-to-local theories, implying that gists are indeed more readily perceived than objects, probably at an earlier stage. Yet the latter finding that objects also influence gist recognition suggests a more parallel and interactive view of both processes than previously assumed.

The self-face captures attention without consciousness: Evidence from the N2pc ERP component analysis.

It is well established that stimuli representing or associated with ourselves, like our own name or an image of our own face, benefit from preferential processing. However, two key questions concerning the self-prioritization mechanism remain to be addressed. First, does it operate in an automatic manner during the early processing, or rather in a more controlled fashion at later processing stages? Second, is it specific to the self-related stimuli, or can it be activated also by other stimuli that are familiar or salient? We conducted a dot-probe experiment to investigate the mechanism behind the attentional prioritization of the self-face image and to tackle both questions. The former, by employing a backwards masking procedure to isolate the early and preconscious processing stages. The latter, by investigating whether a face that becomes visually familiar due to repeated presentations is able to capture attention in a similar manner as the self-face. Analysis of the N2pc ERP component revealed that the self-face image automatically captures attention, both when processed consciously and unconsciously. In contrast, the visually familiar face did not attract attention, neither in the conscious, nor in the unconscious condition. We conclude that the self-prioritization mechanism is early and automatic, and is not triggered by mere visual familiarity. More generally, our results provide further evidence for efficient unconscious processing of faces, and for dissociation between attention and consciousness.

Unconscious perception of one's own name modulates amplitude of the P3B ERP component.

The P3 event-related potential has been known for over 50 years, but its function is still a matter of ongoing debate. Current theories interpret P3b either as a correlate of perceptual consciousness, or as reflecting cognitive processes, like working memory and executive functions. Unexpectedly, recent studies indicate that P3b might occur in response to unconsciously presented stimuli which, if further replicated, will be important for defining its functional role. Therefore, in the present study we investigated the P3b component in response to participants' own name - a stimulus which is subjectively extremely salient and known to evoke a robust P3b response. The self-name and control (other) names were presented supra- and subliminally (backward-masked), in a subjective rating task and in a forced-choice identification task. We found that a consciously perceived self-name evoked a P3b of larger amplitude than the other-name in both tasks, which confirms that the self-name was processed preferentially. When the self-name was presented subliminally it was associated with larger P3b amplitude than the other-name in the identification task, but not in the subjective rating task. This indicates that a salient stimulus can in principle modulate the P3b amplitude even when processed outside of awareness, but also that subliminal processing depends on the task-set and top-down factors. Taken as a whole, our results provide evidence against the interpretation of P3b as a correlate of consciousness (and more generally conflict with the Global Workspace Theory) and will allow a more precise description of the relation between P3b and cognitive processes.

Automatic Attention Capture by Threatening, But Not by Semantically Incongruent Natural Scene Images.

Visual objects are typically perceived as parts of an entire visual scene, and the scene's context provides information crucial in the object recognition process. Fundamental insights into the mechanisms of context-object integration have come from research on semantically incongruent objects, which are defined as objects with a very low probability of occurring in a given context. However, the role of attention in processing of the context-object mismatch remains unclear, with some studies providing evidence in favor, but other against an automatic capture of attention by incongruent objects. Therefore, in the present study, 25 subjects completed a dot-probe task, in which pairs of scenes-congruent and incongruent or neutral and threatening-were presented as task-irrelevant distractors. Importantly, threatening scenes are known to robustly capture attention and thus were included in the present study to provide a context for interpretation of results regarding incongruent scenes. Using N2 posterior-contralateral ERP component as a primary measure, we revealed that threatening images indeed capture attention automatically and rapidly, but semantically incongruent scenes do not benefit from an automatic attentional selection. Thus, our results suggest that identification of the context-object mismatch is not preattentive.

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.

Informativeness of Auditory Stimuli Does Not Affect EEG Signal Diversity.

Brain signal diversity constitutes a robust neuronal marker of the global states of consciousness. It has been demonstrated that, in comparison to the resting wakefulness, signal diversity is lower during unconscious states, and higher during psychedelic states. A plausible interpretation of these findings is that the neuronal diversity corresponds to the diversity of subjective conscious experiences. Therefore, in the present study we varied an information rate processed by the subjects and hypothesized that greater information rate will be related to richer and more differentiated phenomenology and, consequently, to greater signal diversity. To test this hypothesis speech recordings (excerpts from an audio-book) were presented to subjects at five different speeds (65, 83, 100, 117, and 135% of the original speed). By increasing or decreasing speed of the recordings we were able to, respectively, increase or decrease the presented information rate. We also included a backward (unintelligible) speech presentation and a resting-state condition (no auditory stimulation). We tested 19 healthy subjects and analyzed the recorded EEG signal (64 channels) in terms of Lempel-Ziv diversity (LZs). We report the following findings. First, our main hypothesis was not confirmed, as Bayes Factor indicates evidence for no effect when comparing LZs among five presentation speeds. Second, we found that LZs during the resting-state was greater than during processing of both meaningful and unintelligible speech. Third, an additional analysis uncovered a gradual decrease of diversity over the time-course of the experiment, which might reflect a decrease in vigilance. We thus speculate that higher signal diversity during the unconstrained resting-state might be due to a greater variety of experiences, involving spontaneous attention switching and mind wandering.

The impact of self-esteem on the preferential processing of self-related information: Electrophysiological correlates of explicit self vs. other evaluation.

Preferential processing of self-related information is a well-documented phenomenon on both the behavioral and neural levels. However, the impact of self-esteem on this self-preference has not been studied in a systematic way. Here, the electrophysiological correlates of explicit self-reflection were investigated in individuals with low (LSE) and high self-esteem (HSE). Participants evaluated trait adjectives in reference to the self or to an "other" person (close-other, famous) while EEG was recorded. The analysis of event-related potentials focused on the late positive component (LPC), which exhibits a fronto-central distribution and latency over 500 ms. In both LSE and HSE groups, the amplitudes of LPC were enhanced in the self condition when compared to control conditions (both close-other and famous). Crucially, LPC amplitudes in the HSE group were significantly higher than in the LSE group. Moreover, the self-preference effect, defined as the difference between amplitudes of LPC associated with the evaluation of words in relation to oneself vs. other people, was significantly higher in the HSE group than in the LSE group. Overall, our findings indicate that people with high self-esteem tend to engage in self-referential processing to a higher extent.

Exposure to attachment narratives dynamically modulates cortical arousal during the resting state in the listener.

BACKGROUND: Affective stimulation entails changes in brain network patterns at rest, but it is unknown whether exogenous emotional stimulation has a prolonged effect on the temporal dynamics of endogenous cortical arousal. We therefore investigated differences in cortical arousal in the listener following stimulation with different attachment-related narratives. METHODS: Resting-state EEG was recorded from sixteen healthy subjects for ten minutes each with eyes closed: first at baseline and then after passively listening to three affective narratives from strangers about their early childhood experiences (prototypical for insecure-dismissing, insecure-preoccupied, and secure attachment). Using the VIGALL 2.1 algorithm, low or high vigilance stages in consecutive EEG segments were classified, and their dynamic profile was analyzed. Questionnaires assessed the listeners' emotional response to the content of the narrative. RESULTS: As a general effect of preceding affective stimulation, vigilance following the stimulation was significantly elevated compared to baseline rest, and carryover effects in dynamic vigilance profiles were observed. A difference between narrative conditions was revealed for the insecure-dismissing condition, in which the decrease in duration of high vigilance stages was fastest compared to the other two conditions. The behavioral data supported the observation that especially the insecure narratives induced a tendency in the listener to affectively disengage from the narrative content. DISCUSSION: This study revealed carryover effects in endogenous cortical arousal evoked by preceding affective stimulation and provides evidence for attachment-specific dynamic alterations of brain states and individual differences in emotional reactivity.

Loss of consciousness is related to hyper-correlated gamma-band activity in anesthetized macaques and sleeping humans.

Loss of consciousness can result from a wide range of causes, including natural sleep and pharmacologically induced anesthesia. Important insights might thus come from identifying neuronal mechanisms of loss and re-emergence of consciousness independent of a specific manipulation. Therefore, to seek neuronal signatures of loss of consciousness common to sleep and anesthesia we analyzed spontaneous electrophysiological activity recorded in two experiments. First, electrocorticography (ECoG) acquired from 4 macaque monkeys anesthetized with different anesthetic agents (ketamine, medetomidine, propofol) and, second, stereo-electroencephalography (sEEG) from 10 epilepsy patients in different wake-sleep stages (wakefulness, NREM, REM). Specifically, we investigated co-activation patterns among brain areas, defined as correlations between local amplitudes of gamma-band activity. We found that resting wakefulness was associated with intermediate levels of gamma-band coupling, indicating neither complete dependence, nor full independence among brain regions. In contrast, loss of consciousness during NREM sleep and propofol anesthesia was associated with excessively correlated brain activity, as indicated by a robust increase of number and strength of positive correlations. However, such excessively correlated brain signals were not observed during REM sleep, and were present only to a limited extent during ketamine anesthesia. This might be related to the fact that, despite suppression of behavioral responsiveness, REM sleep and ketamine anesthesia often involve presence of dream-like conscious experiences. We conclude that hyper-correlated gamma-band activity might be a signature of loss of consciousness common across various manipulations and independent of behavioral responsiveness.

Breakdown of long-range temporal correlations in brain oscillations during general anesthesia.

Consciousness has been hypothesized to emerge from complex neuronal dynamics, which prevails when brain operates in a critical state. Evidence supporting this hypothesis comes mainly from studies investigating neuronal activity on a short time-scale of seconds. However, a key aspect of criticality is presence of scale-free temporal dependencies occurring across a wide range of time-scales. Indeed, robust long-range temporal correlations (LRTCs) are found in neuronal oscillations during conscious states, but it is not known how LRTCs are affected by loss of consciousness. To further test a relation between critical dynamics and consciousness, we investigated LRTCs in electrocorticography signals recorded from four macaque monkeys during resting wakefulness and general anesthesia induced by various anesthetics (ketamine, medetomidine, or propofol). Detrended Fluctuation Analysis was used to estimate LRTCs in amplitude fluctuations (envelopes) of band-pass filtered signals. We demonstrate two main findings. First, during conscious states all lateral cortical regions are characterized by significant LRTCs of alpha-band activity (7-14 Hz). LRTCs are stronger in the eyes-open than eyes-closed state, but in both states they form a spatial gradient, with anterior brain regions exhibiting stronger LRTCs than posterior regions. Second, we observed a substantial decrease of LRTCs during loss of consciousness, the magnitude of which was associated with the baseline (i.e. pre-anesthesia) state of the brain. Specifically, brain regions characterized by strongest LRTCs during a wakeful baseline exhibited greatest decreases during anesthesia (i.e. "the rich got poorer"), which consequently disturbed the posterior-anterior gradient. Therefore, our results suggest that general anesthesia affects mainly brain areas characterized by strongest LRTCs during wakefulness, which might account for lack of capacities for extensive temporal integration during loss of consciousness.