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.     

From behaviour to consciousness: Translating what animals do to what animals think

Cognitive ethology, an interdisciplinary and comparative branch of zoology, is concerned with the influence of conscious awareness and intention on animal behaviour. It enquires into the evolutionary value of consciousness. However, consciousness is hard to define and any account of animal behaviour based on it will need to take into account both the physical mechanisms that allow for consciousness, and also consider whether we can have knowledge of the phenomenal experience of consciousness in other species. While the first consideration can be investigated scientifically, phenomenal experience needs to be inferred from behaviour, since most animals are not capable of communicating this experience directly. In fact, many accounts of animal behaviour, behavioural ecology in particular, argue that we cannot accurately explain animal behaviour with relation to thoughts or feelings and conscious awareness of them. Rather, we must concern ourselves with what can be objectively observed and measured. Cognitive ethology, however, argues that we cannot give accurate accounts of complex animal behaviour, for example social interactions or tool use, without taking consciousness into account. In this article I will argue that one can justifiably assign and study consciousness in animals through their behaviour, and that an account of certain animals’ behaviour is incomplete without reference to conscious awareness. In other words, behavioural ecology is essentially flawed as it gives, in certain cases, ultimately incorrect accounts of animal behaviour. Firstly it cannot distinguish between behaviour of more and less conscious animals, and secondly, by avoiding any mention of consciousness, it narrows its own scope, and finally cannot explain complex behaviours such as learning in any meaningful way.     

Towards a pluralistic neurobiological understanding of consciousness

Theories of consciousness are often based on the assumption that a single, unified neurobiological account will explain different types of conscious awareness. However, recent findings show that, even within a single modality such as conscious visual perception, the anatomical location, timing, and information flow of neural activity related to conscious awareness vary depending on both external and internal factors. This suggests that the search for generic neural correlates of consciousness may not be fruitful. I argue that consciousness science requires a more pluralistic approach and propose a new framework: joint determinant theory (JDT). This theory may be capable of accommodating different brain circuit mechanisms for conscious contents as varied as percepts, wills, memories, emotions, and thoughts, as well as their integrated experience.     

Singularity and consciousness: A neuropsychological contribution

In common sense experience based on introspection, consciousness is singular. There is only one ‘me’ and that is the one that is conscious. This means that ‘singularity’ is a defining aspect of ‘consciousness’. However, the three main theories of consciousness, Integrated Information, Global Workspace and Recurrent Processing theory, are generally not very clear on this issue. These theories have traditionally relied heavily on neuropsychological observations and have interpreted various disorders, such as anosognosia, neglect and split-brain as impairments in conscious awareness without any reference to ‘the singularity’. In this review, we will re-examine the theoretical implications of these impairments in conscious awareness and propose a new way how to conceptualize consciousness of singularity. We will argue that the subjective feeling of singularity can coexist with several disunified conscious experiences. Singularity awareness may only come into existence due to environmental response constraints. That is, perceptual, language, memory, attentional and motor processes may largely proceed unintegrated in parallel, whereas a sense of unity only arises when organisms need to respond coherently constrained by the affordances of the environment. Next, we examine from this perspective psychiatric disorders and psycho-active drugs. Finally, we present a first attempt to test this hypothesis with a resting state imaging experiment in a split-brain patient. The results suggest that there is substantial coherence of activation across the two hemispheres. These data show that a complete lesioning of the corpus callosum does not, in general, alter the resting state networks of the brain. Thus, we propose that we have separate systems in the brain that generate distributed conscious. The sense of singularity, the experience of a ‘Me-ness’, emerges in the interaction between the world and response-planning systems, and this leads to coherent activation in the different functional networks across the cortex.     

Investigating the Awareness of Remembering

ABSTRACT— There is a marked lack of consensus concerning the best way to learn how conscious experiences arise. In this article, we advocate for scientific approaches that attempt to bring together four types of phenomena and their corresponding theoretical accounts: behavioral acts, cognitive events, neural events, and subjective experience. We propose that the key challenge is to comprehensively specify the relationships among these four facets of the problem of understanding consciousness without excluding any facet. Although other perspectives on consciousness can also be informative, combining these four perspectives could lead to significant progress in explaining a conscious experience such as remembering. We summarize some relevant findings from cognitive neuroscience investigations of the conscious experience of memory retrieval and of memory behaviors that transpire in the absence of the awareness of remembering. These examples illustrate suitable scientific strategies for making progress in understanding consciousness by developing and testing theories that connect the behavioral expression of recall and recognition, the requisite cognitive transactions, the neural events that make remembering possible, and the awareness of remembering.     

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.     

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.     

Dendritic encoding: an alternative to temporal synaptic coding of conscious experience

In this commentary, arguments are made for a dendritic code being preferable to a temporal synaptic code as a model of conscious experience. A temporal firing pattern is a product of an ongoing neural computation; hence, it is based on a neural algorithm and an algorithm may not provide the most suitable model for conscious experience. Reiteration of a temporal firing code as suggested in a preceding article (Helekar, 1999) does not necessarily improve the situation. The alternative model presented here is that certain synaptic activity patterns, possibly those possessing universal features as suggested by Helekar, can become encoded in the dendritic structure. Following dendritic encoding, quantum phenomena in those specific dendrite sets could illuminate the static image of that encoded synaptic activity. It is the activation of the static image that would be equivalent to conscious experience; thus, conscious awareness would not be directly affiliated with synaptic activity. This dendrite encoding model may go farther than other models to explain the gestalt nature of consciousness, insofar as quantum entanglement could produce an interconnectedness between specific sets of dendrites-an interconnectedness that need not be based on neural computation or neural connections.     

Anesthesia, neural information processing, and conscious awareness

Possible systemic effects of general anesthetic agents on neural information processing are discussed in the context of the thalamocortical suppression hypothesis presented by Drs. Alkire, Haier, and Fallon (this issue) in their PET study of the anesthetized state. Accounts of the neural requisites of consciousness fall into two broad categories. Neuronal-specificity theories postulate that activity in particular neural populations is sufficient for conscious awareness, while process-coherence theories postulate that particular organizations of neural activity are sufficient. Accounts of anesthetic narcosis, on the other hand, explain losses of consciousness in terms of neural signal-suppressions, transmission blocks, and the disruptions of signal interpretation. While signal-suppression may account for the actions of some anesthetic agents, the existence of anesthetics, such as choralose, that cause both loss of consciousness and elevated discharge rates, is problematic for a general theory of narcosis that is based purely on signal suppression and transmission-block. However, anesthetic agents also alter relative firing rates and temporal discharge patterns that may disrupt the coherence of neural signals and the functioning of the neural networks that interpret them. It is difficult at present, solely on the basis of regional brain metabolic rates, to test process-coherence hypotheses regarding organizational requisites for conscious awareness. While these pioneering PET studies have great merit as panoramic windows of mind-brain correlates, wider ranges of theory and empirical evidence need to be brought into the formulation of truly comprehensive theories of consciousness and anesthesia.     

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.     

Radical embodiment: neural dynamics and consciousness

We propose a new approach to the neuroscience of consciousness, growing out of the ‘enactive’ viewpoint in cognitive science. This approach aims to map the neural substrates of consciousness at the level of large-scale, emergent and transient dynamical patterns of brain activity (rather than at the level of particular circuits or classes of neurons), and it suggests that the processes crucial for consciousness cut across the brain–body–world divisions, rather than being brain-bound neural events. Whereas standard approaches to the neural correlates of consciousness have assumed a one-way causal-explanatory relationship between internal neural representational systems and the contents of consciousness, our approach allows for theories and hypotheses about the two-way or reciprocal relationship between embodied conscious states and local neuronal activity.     

Demystifying Without Quining: Wittgenstein and Dennett on Qualitative States

Abstract

In his 1991 book Consciousness Explained, Daniel Deimett presents his “Multiple Drafts” model of consciousness. Central to his theory is the rejection of the notion of ‘qualia’ of the existence of the purported ‘qualitative character’ of conscious experience that many argue rules out the possibility of a purely materialist theory of mind. In eliminating qualia from his theory of consciousness, Dennett claims to be following in the footsteps of Wittgenstein, who also had much to say regarding the nature of ‘private’ experience. In this paper I reject this claim and argue that the elimination of qualia plays no part in Wittgenstein’s radical understanding of conscious experience.’¹     

The neural basis of event-time introspection

We explored the neural mechanisms allowing humans to report the subjective onset times of conscious events. Magnetoencephalographic recordings of neural oscillations were obtained while human subjects introspected the timing of sensory, intentional, and motor events during a forced choice task. Brain activity was reconstructed with high spatio-temporal resolution. Event-time introspection was associated with specific neural activity at the time of subjective event onset which was spatially distinct from activity induced by the event itself. Different brain regions were selectively recruited for introspection of different event types, e.g., the bilateral angular gyrus for introspection of intention. Our results suggest that event-time introspection engages specific neural networks to assess the contents of consciousness. Subjective event times should therefore be interpreted as the result of complex interactions between introspection and experience networks, rather than as direct reproduction of the individual’s conscious state or as a mere post hoc interpretation.     

简论里贝特的意识理论

里贝特是人类意识和自由意志的实验研究领域的一个卓越的、先驱性的神经科学家。里贝特的意识研究工作涉及如下四个方面：（1）关于意识研究的认识论原则;（2）对意识现象本性的界定;（3）意识机制的时控理论;（4）对自由意志的阐释和有意识的心智场理论。里贝特的意识研究独树一帜,其时控理论具有坚实可信的实验证据,它从时间维度揭示了有意识的主观体验以及无意识的心智功能与神经活动之间的时间机制。     

New evidence of animal consciousness

This paper reviews evidence that increases the probability that many animals experience at least simple levels of consciousness. First, the search for neural correlates of consciousness has not found any consciousness-producing structure or process that is limited to human brains. Second, appropriate responses to novel challenges for which the animal has not been prepared by genetic programming or previous experience provide suggestive evidence of animal consciousness because such versatility is most effectively organized by conscious thinking. For example, certain types of classical conditioning require awareness of the learned contingency in human subjects, suggesting comparable awareness in similarly conditioned animals. Other significant examples of versatile behavior suggestive of conscious thinking are scrub jays that exhibit all the objective attributes of episodic memory, evidence that monkeys sometimes know what they know, creative tool-making by crows, and recent interpretation of goal-directed behavior of rats as requiring simple nonreflexive consciousness. Third, animal communication often reports subjective experiences. Apes have demonstrated increased ability to use gestures or keyboard symbols to make requests and answer questions; and parrots have refined their ability to use the imitation of human words to ask for things they want and answer moderately complex questions. New data have demonstrated increased flexibility in the gestural communication of swarming honey bees that leads to vitally important group decisions as to which cavity a swarm should select as its new home. Although no single piece of evidence provides absolute proof of consciousness, this accumulation of strongly suggestive evidence increases significantly the likelihood that some animals experience at least simple conscious thoughts and feelings. The next challenge for cognitive ethologists is to investigate for particular animals the content of their awareness and what life is actually like, for them.Donald R. Griffin died on 7 November 2003     

"Binaural rivalry": dichotic listening as a tool for the investigation of the neural correlate of consciousness

Since about two decades neuroscientists have systematically faced the problem of consciousness: the aim is to discover the neural activity specifically related to conscious perceptions, i.e. the biological properties of what philosophers call qualia. In this view, a neural correlate of consciousness (NCC) is a precise pattern of brain activity that specifically accompanies a particular conscious experience. Almost all studies aimed at investigating the NCC have been carried out in the visual system. One of the most promising paradigms is based on sensory stimuli which elicit bistable percepts, as they allow to decouple subjective perception from the characteristics of the physical stimulation. Such kind of perception can be produced in the visual modality by using particular images (e.g. Rubin's vase/face figure) or by presenting two dissimilar stimuli separately to the two eyes (binocular rivalry). The stimuli compete for perceptual dominance and each image is visible in turn for a few seconds, while the other is suppressed. The use of this methodology has led to important findings concerning visual consciousness, which are briefly discussed. For the investigation of auditory consciousness, a similar stimulation paradigm can be achieved by using dichotic listening, consisting in two different stimuli presented each to one ear, which compete for perception (binaural rivalry). The principal aim of the present mini-review is to discuss the few contributes facing the issue of auditory consciousness and to advance the use of dichotic listening and binaural rivalry as valid tools for its investigation.     

Pre-stimulus alpha predicts inattentional blindness

Pre- and post-stimulus oscillatory activity between 8 and 12 hertz, referred to as the alpha-band, correlates with conscious visual awareness of stimuli across a variety of psychophysical tasks. Within an EEG-adapted inattentional blindness task, the current study sought to examine whether this relationship holds for conscious awareness of stimuli under conditions of inattentional blindness. Noticing rates of the task-irrelevant unexpected stimulus were correlated with a significant decrease in alpha power over bilateral parietal-occipital areas during the pre-stimulus interval, and a significant decrease in alpha power over parietal-occipital regions in the right hemisphere during the post-stimulus interval. Findings are taken to imply alpha-band neural activity represents a valid correlate of consciousness that is not confounded by task relevancy or the need for report.     

Can Studies of Autism Teach us About Consciousness of the Physical and the Mental?

Abstract

Most scientists and theorists concerned with the problem of consciousness focus on our consciousness of the physical world (our sensations, feelings, and awareness). In this paper I consider our consciousness of the mental world (our thoughts about thoughts, intentions, wishes, and emotions).The argument is made that these are two distinct forms of consciousness, the evidence for this deriving from studies of autism. Autism is a severe childhood psychiatric condition in which individuals may be conscious of the physical world but not of the mental world. Relevant experimental evidence is described, including some recent neuroimaging studies pointing towards the neural basis of our consciousness of the mental.     

Intentional Action and the Post-Coma Patient

Detecting conscious awareness in a patient emerging from a coma state is problematic, because our standard attributions of conscious awareness rely on interpreting bodily movement as intentional action. Where there is an absence of intentional bodily action, as in the vegetative state, can we reliably assume that there is an absence of conscious awareness? Recent neuroimaging work suggests that we can attribute conscious awareness to some patients in a vegetative state by interpreting their brain activity as intentional mental action. I suggest that this change of focus, from the interpretation of motor behaviour as intentional bodily action to the interpretation of neural activity as intentional mental action, raises philosophical issues that affect the interpretation of the neuroimaging data.     

Consciousness or the Physical Universe – Which Came First?

Historically, many have seen the intelligibility of the physical universe as showing that it is somehow ultimately dependent upon conscious intelligent pre‐existing being – ‘God’. Today, however, many believe that modern advances in our scientific understanding of the origins and nature of the universe, and of the conscious intelligent beings it contains, render God, as Laplace said, an ‘unnecessary hypothesis’. This article considers whether the findings of modern science do indeed diminish the plausibility of belief in a creator God. Or, on the contrary, whether there are features of current scientific understanding which may reasonably be thought to support the belief that conscious intelligent being pre‐existed the physical universe and caused it to be. In short: can science reasonably be thought to support the view that consciousness created the physical universe rather than that the physical universe created consciousness?