Neural correlates of consciousness reconsidered

It is widely accepted among philosophers that neuroscientists are conducting a search for the neural correlates of consciousness, or NCC. Chalmers (2000) conceptualized this research program as the attempt to correlate the contents of conscious experience with the contents of representations in specific neural populations. A notable claim on behalf of this interpretation is that the neutral language of "correlates" frees us from philosophical disputes over the mind/body relation, allowing the science to move independently. But the experimental paradigms and explanatory canons of neuroscience are not neutral about the mechanical relation between consciousness and the brain. I argue that NCC research is best characterized as an attempt to locate a causally relevant neural mechanism and not as an effort to identify a discrete neural representation, the content of which correlates with some actual experience. It might be said that the first C in "NCC" should stand for "causes" rather than "correlates."     

A field theory of consciousness.

This article summarizes a variety of current as well as previous research in support of a new theory of consciousness. Evidence has been steadily accumulating that information about a stimulus complex is distributed to many neuronal populations dispersed throughout the brain and is represented by the departure from randomness of the temporal pattern of neural discharges within these large ensembles. Zero phase lag synchronization occurs between discharges of neurons in different brain regions and is enhanced by presentation of stimuli. This evidence further suggests that spatiotemporal patterns of coherence, which have been identified by spatial principal component analysis, may encode a multidimensional representation of a present or past event. How such distributed information is integrated into a holistic precept constitutes the binding problem. How a precept defined by a spatial distribution of nonrandomness can be subjectively experienced constitutes the problem of consciousness. Explanations based on a discrete connectionistic network cannot be reconciled with the relevant facts. Evidence is presented herein of invariant features of brain electrical activity found to change reversibly with loss and return of consciousness in a study of 176 patients anesthetized during surgical procedures. A review of relevant research areas, as well as the anesthesia data, leads to a postulation that consciousness is a property of quantum-like processes, within a brain field resonating within a core of structures, which may be the neural substrate of consciousness. This core includes regions of the prefrontal cortex, the frontal cortex, the pre- and paracentral cortex, thalamus, limbic system, and basal ganglia.     

A cross-order integration hypothesis for the neural correlate of consciousness

One major problem many hypotheses regarding the neural correlate of consciousness (NCC), face is what we might call "the why question": why would this particular neural feature, rather than another, correlate with consciousness? The purpose of the present paper is to develop an NCC hypothesis that answers this question. The proposed hypothesis is inspired by the cross-order integration (COI) theory of consciousness, according to which consciousness arises from the functional integration of a first-order representation of an external stimulus and a second-order representation of that first-order representation. The proposal comes in two steps. The first step concerns the "general shape" of the NCC and can be directly derived from COI theory. The second step is a concrete hypothesis that can be arrived at by combining the general shape with empirical considerations.     

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.     

The thalamic dynamic core theory of conscious experience

I propose that primary conscious awareness arises from synchronized activity in dendrites of neurons in dorsal thalamic nuclei, mediated particularly by inhibitory interactions with thalamic reticular neurons. In support, I offer four evidential pillars: (1) consciousness is restricted to the results of cortical computations; (2) thalamus is the common locus of action of brain injury in vegetative state and of general anesthetics; (3) the anatomy and physiology of the thalamus imply a central role in consciousness; (4) neural synchronization is a neural correlate of consciousness.     

`Funda-Mentality': is the conscious mind subtly linked to a basic level of the universe?

Age-old battle lines over the puzzling nature of mental experience are shaping a modern resurgence in the study of consciousness. On one side are the long-dominant `physicalists' who view consciousness as an emergent property of the brain's neural networks. On the alternative, rebellious side are those who see a necessary added ingredient: proto-conscious experience intrinsic to reality, perhaps understandable through modern physics (panpsychists, pan-experientialists, `funda-mentalists'). It is argued here that the physicalist premise alone is unable to solve completely the difficult issues of consciousness and that to do so will require supplemental panpsychist/pan-experiential philosophy expressed in modern physics. In one scheme proto-conscious experience is a basic property of physical reality accessible to a quantum process associated with brain activity. The proposed process is Roger Penrose's `objective reduction' (OR), a self-organizing `collapse' of the quantum wave function related to instability at the most basic level of space-time geometry. In the Penrose-Hameroff model of `orchestrated objective reduction' (Orch OR), OR quantum computation occurs in cytoskeletal microtubules within the brain's neurons. The basic thesis is that consciousness involves brain activities coupled to self-organizing ripples in fundamental reality.     

Missing piece of the puzzle in the science of consciousness: Resting state and endogenous correlates of consciousness

Consciousness still stands as one of the most interesting and the most elusive problems of neuroscience. Finding its correlates is the first step toward its satisfactory explanation. Several theories have proposed its correlates but none of them seem to be generally accepted even though most of them share some very similar elements. These elements are the activity of the thalamus, which is considered by some as the central region for consciousness, and gamma synchronization, which should be the general principal for the emergence of conscious experience. However, all of these proposed theories share one characteristic and that is that they do not take into consideration the recently discovered endogenous activity of the brain, which is generally associated with the default mode network. Although the activity of this large scale brain network is in correlation with various levels of consciousness it is still missing in discussions of consciousness. This review recognizes the importance of endogenous activity and points out the important discoveries of endogenous activity that could be an important step toward a satisfactory explanation of consciousness.     

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.     

A sensorimotor account of vision and visual consciousness

Many current neurophysiological, psychophysical, and psychological approaches to vision rest on the idea that when we see, the brain produces an internal representation of the world. The activation of this internal representation is assumed to give rise to the experience of seeing. The problem with this kind of approach is that it leaves unexplained how the existence of such a detailed internal representation might produce visual consciousness. An alternative proposal is made here. We propose that seeing is a way of acting. It is a particular way of exploring the environment. Activity in internal representations does not generate the experience of seeing. The outside world serves as its own, external, representation. The experience of seeing occurs when the organism masters what we call the governing laws of sensorimotor contingency. The advantage of this approach is that it provides a natural and principled way of accounting for visual consciousness, and for the differences in the perceived quality of sensory experience in the different sensory modalities. Several lines of empirical evidence are brought forward in support of the theory, in particular: evidence from experiments in sensorimotor adaptation, visual "filling in," visual stability despite eye movements, change blindness, sensory substitution, and color perception.     

No representation without awareness in the lateral occipital cortex

Recent research has shown that four small dots presented in the vicinity of, but not adjacent to, a target stimulus can banish that stimulus from conscious awareness. It is thought that the mental representation of the masked stimulus is "erased" by the trailing quartet of dots. Using functional magnetic resonance adaptation, we show that there is no persisting neural representation of the successfully masked stimulus in lateral occipital cortex, a region that has been implicated in the processing of object structure. This finding rules out the alternative interpretation that a lingering neural representation is merely rendered inaccessible to consciousness, as is the fate, for example, of monocular information under conditions of binocular rivalry.     

Synchronous neural oscillations and cognitive processes

The central problem for cognitive neuroscience is to describe how cognitive processes arise from brain processes. This review summarizes the recent evidence that synchronous neural oscillations reveal much about the origin and nature of cognitive processes such as memory, attention and consciousness. Memory processes are most closely related to theta and gamma rhythms, whereas attention seems closely associated with alpha and gamma rhythms. Conscious awareness may arise from synchronous neural oscillations occurring globally throughout the brain rather than from the locally synchronous oscillations that occur when a sensory area encodes a stimulus. These associations between the dynamics of the brain and cognitive processes indicate progress towards a unified theory of brain and cognition.     

A global workspace model for phenomenal and access consciousness

Both the global workspace theory and Block’s distinction between phenomenal and access consciousness, are central in the current debates about consciousness and the neural correlates of consciousness. In this article, a unifying global workspace model for phenomenal and access consciousness is proposed. In the model, recurrent neural interactions take place in distinct yet interacting access and phenomenal brain loops. The effectiveness of feedback signaling onto sensory cortical maps is emphasized for the neural correlates of phenomenal consciousness. Two forms of top-down attention, attention for perception and attention for access, play differential roles for phenomenal and access consciousness. The model is implemented in a neural network form, with the simulation of single and multiple visual object processing, and of the attentional blink.     

Predictive minds and small-scale models: Kenneth Craik’s contribution to cognitive science

I identify three lessons from Kenneth Craik’s landmark book “The Nature of Explanation” for contemporary debates surrounding the existence, extent, and nature of mental representation: first, an account of mental representations as neural structures that function analogously to public models; second, an appreciation of prediction as the central component of intelligence in demand of such models; and third, a metaphor for understanding the brain as an engineer, not a scientist. I then relate these insights to discussions surrounding the representational status of predictive processing – which, I argue, provides a contemporary vindication of Craik’s extremely prescient “hypothesis on the nature of thought.”     

Are we explaining consciousness yet?

Theorists are converging from quite different quarters on a version of the global neuronal workspace model of consciousness, but there are residual confusions to be dissolved. In particular, theorists must resist the temptation to see global accessibility as the cause of consciousness (as if consciousness were some other, further condition); rather, it is consciousness. A useful metaphor for keeping this elusive idea in focus is that consciousness is rather like fame in the brain. It is not a privileged medium of representation, or an added property some states have; it is the very mutual accessibility that gives some informational states the powers that come with a subject's consciousness of that information. Like fame, consciousness is not a momentary condition, or a purely dispositional state, but rather a matter of actual influence over time. Theorists who take on the task of accounting for the aftermath that is critical for consciousness often appear to be leaving out the Subject of consciousness, when in fact they are providing an analysis of the Subject, a necessary component in any serious theory of consciousness.     

A connectionist theory of phenomenal experience

When cognitive scientists apply computational theory to the problem of phenomenal consciousness, as many have been doing recently, there are two fundamentally distinct approaches available. Consciousness is to be explained either in terms of the nature of the representational vehicles the brain deploys or in terms of the computational processes defined over these vehicles. We call versions of these two approaches vehicle and process theories of consciousness, respectively. However, although there may be space for vehicle theories of consciousness in cognitive science, they are relatively rare. This is because of the influence exerted, on the one hand, by a large body of research that purports to show that the explicit representation of information in the brain and conscious experience are dissociable, and on the other, by the classical computational theory of mind--the theory that takes human cognition to be a species of symbol manipulation. Two recent developments in cognitive science combine to suggest that a reappraisal of this situation is in order. First, a number of theorists have recently been highly critical of the experimental methodologies used in the dissociation studies--so critical, in fact, that it is no longer reasonable to assume that the dissociability of conscious experience and explicit representation has been adequately demonstrated. Second, classicism, as a theory of human cognition, is no longer as dominant in cognitive science as it once was. It now has a lively competitor in the form of connectionism; and connectionism, unlike classicism, does have the computational resources to support a robust vehicle theory of consciousness. In this target article we develop and defend this connectionist vehicle theory of consciousness. It takes the form of the following simple empirical hypothesis: phenomenal experience consists of the explicit representation of information in neurally realized parallel distributed processing (PDP) networks. This hypothesis leads us to reassess some common wisdom about consciousness, but, we argue, in fruitful and ultimately plausible ways.     

The precuneus and consciousness

This article reviews the rapidly growing literature on the functional anatomy and behavioral correlates of the precuneus, with special reference to imaging neuroscience studies using hamodynamic techniques. The precuneus, along with adjacent areas within the posteromedial parietal cortex, is among the most active cortical regions according to the "default mode" of brain function during the conscious resting state, whereas it selectively deactivates in a number of pathophysiological conditions (ie, sleep, vegetative state, drug-induced anesthesia), and neuropsychiatric disorders (ie, epilepsy, Alzheimer's disease, and schizophrenia) characterized by impaired consciousness. These findings, along with the widespread connectivity pattern, suggest that the precuneus may play a central role in the neural network correlates of consciousness. Specifically, its activity seems to correlate with self-reflection processes, possibly involving mental imagery and episodic/autobiographical memory retrieval.     

Affective consciousness: Core emotional feelings in animals and humans

The position advanced in this paper is that the bedrock of emotional feelings is contained within the evolved emotional action apparatus of mammalian brains. This dual-aspect monism approach to brain-mind functions, which asserts that emotional feelings may reflect the neurodynamics of brain systems that generate instinctual emotional behaviors, saves us from various conceptual conundrums. In coarse form, primary process affective consciousness seems to be fundamentally an unconditional "gift of nature" rather than an acquired skill, even though those systems facilitate skill acquisition via various felt reinforcements. Affective consciousness, being a comparatively intrinsic function of the brain, shared homologously by all mammalian species, should be the easiest variant of consciousness to study in animals. This is not to deny that some secondary processes (e.g., awareness of feelings in the generation of behavioral choices) cannot be evaluated in animals with sufficiently clever behavioral learning procedures, as with place-preference procedures and the analysis of changes in learned behaviors after one has induced re-valuation of incentives. Rather, the claim is that a direct neuroscientific study of primary process emotional/affective states is best achieved through the study of the intrinsic ("instinctual"), albeit experientially refined, emotional action tendencies of other animals. In this view, core emotional feelings may reflect the neurodynamic attractor landscapes of a variety of extended trans-diencephalic, limbic emotional action systems-including SEEKING, FEAR, RAGE, LUST, CARE, PANIC, and PLAY. Through a study of these brain systems, the neural infrastructure of human and animal affective consciousness may be revealed. Emotional feelings are instantiated in large-scale neurodynamics that can be most effectively monitored via the ethological analysis of emotional action tendencies and the accompanying brain neurochemical/electrical changes. The intrinsic coherence of such emotional responses is demonstrated by the fact that they can be provoked by electrical and chemical stimulation of specific brain zones-effects that are affectively laden. For substantive progress in this emerging research arena, animal brain researchers need to discuss affective brain functions more openly. Secondary awareness processes, because of their more conditional, contextually situated nature, are more difficult to understand in any neuroscientific detail. In other words, the information-processing brain functions, critical for cognitive consciousness, are harder to study in other animals than the more homologous emotional/motivational affective state functions of the brain.     

Afterimages: a tool for defining the neural correlate of visual consciousness

Our visual system not only mediates information about the visual environment but is capable of generating pictures of nonexistent worlds: afterimages, illusions, phosphenes, etc. We are "aware" of these pictures just as we are aware of the images of natural, physical objects. This raises the question: is the neural correlate of consciousness (NCC) of such images the same as that of images of physical objects? Images of natural objects have some properties in common with afterimages (e.g., stability of verticality) but there are also obvious differences (e.g., images maintain size constancy, whereas afterimages follow Emmert's Law: when seen while screens at different distances are observed, an afterimage looks larger, the greater the distance of the screen). The differences can be explained by differences in the retinal extent of images and afterimages, which favors the view that both have the same NCC. It seems reasonable to assume that before neural activity can produce awareness, all the computations necessary for a veridical representation of, e.g., an object, must be completed within the neural substrate and that information characteristic of a particular object must be available within the NCC. Given these assumptions, it can be shown that no retinotopic (in a strict sense) cortical areas can serve as the NCC, although some type of topographic representation is necessary. It seems also to be unlikely that neurons classified as cardinal cells alone can serve as NCC.