The earliest electrophysiological correlate of visual awareness?

To examine the neural correlates and timing of human visual awareness, we recorded event-related potentials (ERPs) in two experiments while the observers were detecting a grey dot that was presented near subjective threshold. ERPs were averaged for conscious detections of the stimulus (hits) and nondetections (misses) separately. Our results revealed that hits, as compared to misses, showed a negativity around 180-350 ms at occipital and posterior temporal sites. It was followed by a positive wave after 400-500 ms, peaking at parietal sites. These correlates were not affected by a manipulation of attention. The early negativity, called 'visual awareness negativity' (VAN), may be a general, primary electrophysiological correlate of visual awareness. The present data show that it can be observed in response to appearance of a stimulus in visual awareness and that it generalizes across different manipulations of stimulus visibility.     

Neurological disorders and the structure of human consciousness

Recent studies that identify distinct neural correlates of perceptual awareness offer a promising step towards improved understanding of the neurological underpinnings of conscious experience. Such studies indicate that perceptual awareness is modular in nature, with neural correlates of awareness consisting of the specialized structures involved in perceptual processing. However, the integrative, multimodal nature of conscious experience appears to require a functional architecture that overcomes this modular segregation of function. We propose a model in which experience emerges from the dynamic interactions of specialized component processes via a distributed neural network. Such a model offers a mechanism to explain several empirical observations of the neural correlates of perceptual awareness, cognitive function, and symptoms of neurological damage.     

Neural events and perceptual awareness

Neural correlates of perceptual awareness, until very recently an elusive quarry, are now almost commonplace findings. This article first describes a variety of neural correlates of perceptual awareness based on fMRI, ERPs, and single-unit recordings. It is then argued that our quest should ultimately focus not on mere correlates of awareness, but rather on the neural events that are both necessary and sufficient for perceptual awareness. Indeed, preliminary evidence suggests that although many of the neural correlates already reported may be necessary for the corresponding state of awareness, it is unlikely that they are sufficient for it. The final section considers three hypotheses concerning the possible sufficiency conditions for perceptual awareness.     

Psychophysical magic: rendering the visible 'invisible'

What are the neural correlates of conscious visual awareness? Tackling this question requires contrasting neural correlates of stimulus processing culminating in visual awareness with neural correlates of stimulus processing unaccompanied by awareness. To produce these two neural states, one must be able to erase an otherwise visible stimulus from awareness. This article describes and assesses visual phenomena involving dissociation of physical stimulation and conscious awareness: degraded stimulation, visual masking, visual crowding, bistable figures, binocular rivalry, motion-induced blindness, inattentional blindness, change blindness and attentional blink. No single approach stands above the others, but those producing changing visual awareness despite invariant physical stimulation are clearly preferable. Such phenomena can help lead us ultimately to a comprehensive account of the neural correlates of conscious awareness.     

On the possibility of universal neural coding of subjective experience

Various neurophysiological experiments have revealed remarkable correlations between cortical neuronal activity and subjective experiences. However, the mere presence of neuronal electrical activity does not appear to be sufficient to produce these experiences. It has been suggested that the explanation for the neural basis of consciousness might lie in understanding the reason that some types of neuronal activity possess subjective correlates and others do not. Here I propose and develop the idea that this difference may be caused by the existence of an elementary nonarbitrary linkage between temporal or spatiotemporal patterns of neuronal activity and their subjective attributes. I also show how cortical neural circuits capable of generating experience-coding patterns could emerge during evolution and brain development, due to the presence of spontaneous stochastic neuronal activity and activity-dependent synaptic plasticity. This hypothesis leads to several testable predictions, principal among which is the idea that the neural correlates of consciousness are essentially innate and universal.     

In and out of consciousness: Sustained electrophysiological activity reflects individual differences in perceptual awareness

Although significant advances in our understanding of the cognitive and neural processes involved in conscious awareness have occurred in recent years, the precise mechanisms that support consciousness remain elusive. Examining the neural correlates associated with the moment a stimulus enters or exits conscious awareness is one way to potentially identify the neural mechanisms that give rise to consciousness. In the present study, we recorded neural activity using electroencephalography (EEG) while participants observed a bilateral shape-from-motion (SFM) display. While the display is in motion, the observer perceives an object that is immediately segregated from a noisy background. After the motion stops, the observer's experience of the object remains momentarily in awareness, before it eventually fades out of consciousness back into the noisy background. Consistent with subjective reports of perceptual experience, we observed a prominent sustained posterior contralateral negativity known as the contralateral delay activity (CDA). This activity was sustained only in conditions associated with sustained awareness. Interestingly, the amplitude of the CDA was correlated with individual differences in visual awareness, suggesting that this activity plays a significant role in the maintenance of objects in consciousness. The CDA is typically associated with visual short-term memory (VSTM), suggesting that conscious visual awareness may be mediated by the same neural and cognitive mechanisms that support VSTM. Our results demonstrate that the CDA may reflect the contents of conscious awareness, and therefore can provide a measure to track when information moves in and out of consciousness.     

In defense of experience-coding nonarbitrary temporal neural activity patterns

Various neurophysiological experiments have revealed remarkable correlations between cortical neuronal activity and subjective experiences. However, the mere presence of neuronal electrical activity does not appear to be sufficient to produce these experiences. It has been suggested that the explanation for the neural basis of consciousness might lie in understanding the reason that some types of neuronal activity possess subjective correlates and others do not. Here I propose and develop the idea that this difference may be caused by the existence of an elementary nonarbitrary linkage between temporal or spatiotemporal patterns of neuronal activity and their subjective attributes. I also show how cortical neural circuits capable of generating experience-coding patterns could emerge during evolution and brain development, due to the presence of spontaneous stochastic neuronal activity and activity-dependent synaptic plasticity. This hypothesis leads to several testable predictions, principal among which is the idea that the neural correlates of consciousness are essentially innate and universal.     

Neural synchronization as a hypothetical explanation of the psychoanalytic unconscious

Cognitive scientists have tried to explain the neural mechanisms of unconscious mental states such as coma, epileptic seizures, and anesthesia-induced unconsciousness. However these types of unconscious states are different from the psychoanalytic unconscious. In this review, we aim to present our hypothesis about the neural correlates underlying psychoanalytic unconscious. To fulfill this aim, we firstly review the previous explanations about the neural correlates of conscious and unconscious mental states, such as brain oscillations, synchronicity of neural networks, and cognitive binding. By doing so, we hope to lay a neuroscientific ground for our hypothesis about neural correlates of psychoanalytic unconscious; parallel but unsynchronized neural networks between different layers of consciousness and unconsciousness. Next, we propose a neuroscientific mechanism about how the repressed mental events reach the conscious awareness; the lock of neural synchronization between two mental layers of conscious and unconscious. At the last section, we will discuss the data about schizophrenia as a clinical example of our proposed hypothesis.     

Identifying neural correlates of consciousness: the state space approach

This article sketches an idealized strategy for the identification of neural correlates of consciousness. The proposed strategy is based on a state space approach originating from the analysis of dynamical systems. The article then focuses on one constituent of consciousness, phenomenal awareness. Several rudimentary requirements for the identification of neural correlates of phenomenal awareness are suggested. These requirements are related to empirical data on selective attention, on completely intrinsic selection and on globally unconscious states. As an example, neuroscientific findings on synchronized gamma activity are categorized according to these requirements.     

Are errors detected before they occur? Early error sensations revealed by metacognitive judgments on the timing of error awareness

Errors in choice tasks are not only detected fast and reliably, participants often report that they knew that an error occurred already before a response was produced. These early error sensations stand in contrast with evidence suggesting that the earliest neural correlates of error awareness emerge around 300 ms after erroneous responses. The present study aimed to investigate whether anecdotal evidence for early error sensations can be corroborated in a controlled study in which participants provide metacognitive judgments on the subjective timing of error awareness. In Experiment 1, participants had to report whether they became aware of their errors before or after the response. In Experiment 2, we measured confidence in these metacognitive judgments. Our data show that participants report early error sensations with high confidence in the majority of error trials across paradigms and experiments. These results provide first evidence for early error sensations, informing theories of error awareness.     

The neural correlates of conscious experience: an experimental framework

Demonstrating that neural activity ‘represents’ physical properties of the world such as the orientation of a line in the receptive field of a nerve cell is a standard procedure in neuroscience. However, not all such neural activity will be associated with the mental representations that form the contents of consciousness. In some cases, such as when patients with blindsight correctly ‘guess’ the location of a stimulus, neural activity is associated with physical stimulation and with appropriate behaviour, but not with awareness. To identify the neural correlates of conscious experience we need to identify patterns of neural activity that are specifically associated with awareness. Experiments aimed at making such identifications require that subjects report some aspect of their conscious experience either verbally or through some pre-arranged non-verbal report while neural activity is measured. If there is some characteristic neural signature of consciousness, then this should be distinguishable from the kinds of neural activity associated with stimulation and/or behaviour in the absence of awareness. It remains to be seen whether the neural signature of consciousness relates to the location of the neural activity, the temporal properties of the neural activity or the form of the interaction between activity in different brain regions.     

The ‘I’ and the ‘Me’ in self-referential awareness: a neurocognitive hypothesis

The nature of the ‘self’ and self-referential awareness has been one of the most debated issues in philosophy, psychology and cognitive neuroscience. Understanding the neurocognitive bases of self-related representation and processing is also crucial to research on the neural correlates of consciousness. The distinction between an ‘I’, corresponding to a subjective sense of the self as a thinker and causal agent, and a ‘Me’, as the objective sense of the self with the unique and identifiable features constituting one’s self-image or self-concept, suggested by William James, has been re-elaborated by authors from different theoretical perspectives. In this article, empirical studies and theories about the ‘I’ and the ‘Me’ in cognition and self-related awareness are reviewed, including the relationships between self and perception, self and memory, the development of the self, self-referential stimulus processing, as well as related neuroimaging studies. Subsequently, the relations between self and different aspects of consciousness are considered. On the basis of the reviewed literature and with reference to Block’s distinction between phenomenal and access consciousness, a neurocognitive hypothesis is formulated about ‘I’-related and ‘Me’-related self-referential awareness. This hypothesis is extended to metacognitive awareness and a form of non-transitive consciousness, characteristic of meditation experiences and studies, with particular reference to the notion of mindfulness and other Buddhist constructs.     

情绪面孔的意识神经相关物及其无意识自动加工：来自事件相关电位的证据

意识的神经相关物尚有争议, 且个体能否无意识自动检测视觉环境变化尚不清楚。本研究采用非注意视盲范式操控视觉意识, 并引入具有社会信息的情绪面孔, 探讨意识的神经相关物以及视觉意识与自动检测变化机制的关系。在A阶段, 部分被试对任务无关的情绪面孔处于无意识水平; 在B阶段, 所有被试对任务无关的情绪面孔处于意识水平; 在C阶段, 所有被试对任务相关的情绪面孔处于意识水平。结果显示, 任务无关的情绪面孔的意识过程诱发视觉意识负波(visual awareness negativity, VAN)、晚期正成分(late positivity, LP)和晚期枕区正成分(late occipital positivity, LOP)。此外, 无意识的情绪面孔能诱发视觉失匹配负波(visual mismatch negativity, vMMN), 且其幅值不受意识影响, 但是受任务相关性调制。这些结果提示对情绪面孔的视觉意识在不同的时间进程上有不同的ERP指标——VAN反映早期知觉经验, 而LP和LOP反映晚期意识过程, 而且面孔情绪信息的自动加工独立于视觉意识, 但是受视觉注意调制。     

Visual awareness negativity is an early neural correlate of awareness: A preregistered study with two Gabor sizes

Electrophysiological recordings are commonly used to study the neural correlates of consciousness in humans. Previous research is inconsistent as to whether awareness can be indexed with visual awareness negativity (VAN) at about 200 ms or if it occurs later. The present study was preregistered with two main aims: First, to provide independent evidence for or against the presence of VAN, and second, to study whether stimulus size may account for the inconsistent findings. Subjects were shown low-contrast Gaussian filtered gratings (Gabor patches) in the four visual quadrants. Gabor size (large and small) was varied in different sessions and calibrated to each subject’s threshold of visual awareness. Event-related potentials were derived from trials in which subjects localized the Gabors correctly to capture the difference between trials in which they reported awareness versus no awareness. Bayesian analyses revealed very strong evidence for the presence of VAN for both Gabor sizes. However, there was no evidence for or against an effect of stimulus size. The present findings provide evidence for VAN as an early neural correlate of awareness.     

‘Eppur si move’: The Association Between Electrophysiological and Psychophysical Signatures of Perceived Movement Illusions

Vibration applied to muscle tendons induces an illusory perception of movement by activating spindle receptors and Ia afferent fibers. While neural structures supporting such illusions are quite defined, functional and psychophysical correlates of their perception have not been deeply investigated. The authors aimed at exploring subjective perception of illusory movements thanks to a repeated-stimulation multimodal approach. The integrated analysis of electroencephalography, source data (standardized low-resolution brain electromagnetic tomography–sLORETA), and psychophysical evaluation pointed at the activation of a frontoparietal sensorimotor network, with prefrontal-medial regions mainly involved in supporting the awareness for illusory movements. Further, illusions showed consistency in terms of presence, vividness, and duration. Finally, superior parietal structures seemed to play an important role in self-attribution of agency and ownership during such experiences.     

Measuring consciousness: Is one measure better than the other?

What is the best way of assessing the extent to which people are aware of a stimulus? Here, using a masked visual identification task, we compared three measures of subjective awareness: The Perceptual Awareness Scale (PAS), through which participants are asked to rate the clarity of their visual experience; confidence ratings (CR), through which participants express their confidence in their identification decisions, and Post-decision wagering (PDW), in which participants place a monetary wager on their decisions. We conducted detailed explorations of the relationships between awareness and identification performance, looking to determine (1) which scale best correlates with performance, and (2) whether we can detect performance in the absence of awareness and how the scales differ from each other in terms of revealing such unconscious processing. Based on these findings we discuss whether perceptual awareness should be considered graded or dichotomous. Results showed that PAS showed a much stronger performance-awareness correlation than either CR or PDW, particularly for low stimulus intensities. In general, all scales indicated above-chance performance when participants claimed not to have seen anything. However, such above-chance performance only showed when we also observed a correlation between awareness and performance. Thus (1) PAS seems to be the most exhaustive measure of awareness, and (2) we find support for above-chance performance in the absence of subjective awareness, but such unconscious knowledge only contributes to performance when we observe conscious knowledge as well. Similarities and differences between scales are discussed in the light of consciousness theories and response strategies.     

Generalization of skills between operant control and discrimination of EEG alpha

While biofeedback is often said to increase self-control of physiological states by increasing awareness of their subjective correlates, relatively few studies have analyzed the relationship between control (standard biofeedback) and awareness (a discrimination paradigm). We hypothesized that the two skills would generalize and facilitate each other for 8–12 Hz EEG amplitude (alpha). Participants were given 7 sessions of training to either control or discriminate Pz alpha followed by 3 sessions of the other paradigm. Another group was given 7 sessions with time divided equally between the two types of training. The control-training first group showed significant generalization of skills to the discrimination task. However, the reverse was not true, and the combined task group did no better in either task than the other two groups. These results provide ambivalent support for the role of awareness in biofeedback, and suggest possible improvements in the discrimination paradigm.     

Implicit threat learning involves the dorsolateral prefrontal cortex and the cerebellum

Background/ObjectiveMost studies investigating the neural correlates of threat learning were carried out using an explicit Pavlovian conditioning paradigm where declarative knowledge on contingencies between conditioned (CS) and unconditioned stimuli (US) is acquired. The current study aimed at understanding the neural correlates of threat conditioning when contingency awareness is limited or even absent.MethodWe conducted an fMRI report of threat learning in an implicit associative learning paradigm called multi-CS conditioning, in which a number of faces were associated with aversive screams (US) such that participants could not report contingencies between the faces and the screams.ResultsThe univariate results showed support for the recruitment of threat-related regions including the dorsolateral prefrontal cortex (dlPFC) and the cerebellum during acquisition. Further analyses by the multivariate representational similarity technique identified learning-dependent changes in the bilateral dlPFC.ConclusionOur findings support the involvement of the dlPFC and the cerebellum in threat conditioning that occurs with highly limited or even absent contingency awareness.     

The neural correlate of (un)awareness: lessons from the vegetative state

Consciousness has two main components: wakefulness and awareness. The vegetative state is characterized by wakefulness without awareness. Recent functional neuroimaging results have shown that some parts of the cortex are still functioning in 'vegetative' patients. External stimulation, such as a painful stimulus, still activates 'primary' sensory cortices in these patients but these areas are functionally disconnected from 'higher order' associative areas needed for awareness. Such studies are disentangling the neural correlates of the vegetative state from the minimally conscious state, and have major clinical consequences in addition to empirical importance for the understanding of consciousness.