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.     

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."     

Two neural correlates of consciousness

Neuroscientists continue to search for "the" neural correlate of consciousness (NCC). In this article, I argue that a framework in which there are at least two distinct NCCs is increasingly making more sense of empirical results than one in which there is a single NCC. I outline the distinction between phenomenal NCC and access NCC, and show how they can be distinguished by experimental approaches, in particular signal-detection theory approaches. Recent findings in cognitive neuroscience provide an empirical case for two different NCCs.     

“Paradox of slow frequencies” – Are slow frequencies in upper cortical layers a neural predisposition of the level/state of consciousness (NPC)?

Consciousness research has much focused on faster frequencies like alpha or gamma while neglecting the slower ones in the infraslow (0.001–0.1 Hz) and slow (0.1–1 Hz) frequency range. These slower frequency ranges have a “bad reputation” though; their increase in power can observed during the loss of consciousness as in sleep, anesthesia, and vegetative state. However, at the same time, slower frequencies have been conceived instrumental for consciousness. The present paper aims to resolve this paradox which I describe as “paradox of slow frequencies”. I first show various data that suggest a central role of slower frequencies in integrating faster ones, i.e., “temporo-spatial integration and nestedness”. Such “temporo-spatial integration and nestedness” is disrupted during the loss of consciousness as in anesthesia and sleep leading to “temporo-spatial fragmentation and isolation” between slow and fast frequencies. Slow frequencies are supposedly mediated by neural activity in upper cortical layers in higher-order associative regions as distinguished from lower cortical layers that are related to faster frequencies. Taken together, slower and faster frequencies take on different roles for the level/state of consciousness. Faster frequencies by themselves are sufficient and thus a neural correlate of consciousness (NCC) while slower frequencies are a necessary non-sufficient condition of possible consciousness, e.g., a neural predisposition of the level/state of consciousness (NPC). This resolves the “paradox of slow frequencies” in that it assigns different roles to slower and faster frequencies in consciousness, i.e., NCC and NPC. Taken as NCC and NPC, fast and slow frequencies including their relation as in “temporo-spatial integration and nestedness” can be considered a first “building bloc” of a future “temporo-spatial theory of consciousness” (TTC) (Northoff, 2013; Northoff, 2014b; Northoff & Huang, 2017).     

Physical,neural, and mental timing

The conclusions drawn by Benjamin Libet from his work with colleagues on the timing of somatosensorial conscious experiences has met with a lot of praise and criticism. In this issue we find three examples of the latter. Here I attempt to place the divide between the two opponent camps in a broader perspective by analyzing the question of the relation between physical timing, neural timing, and experiential (mental) timing. The nervous system does a sophisticated job of recombining and recoding messages from the sensorial surfaces and if these processes are slighted in a theory, it might become necessary to postulate weird operations, including subjective back-referral. Neuroscientifically inspired theories are of necessity still based on guesses, extrapolations, and philosophically dubious manners of speech. They often assume some neural correlate of consciousness (NCC) as a part of the nervous system that transforms neural activity in reportable experiences. The majority of neuroscientists appear to assume that the NCC can compare and bind activity patterns only if they arrive simultaneously at the NCC. This leads to a search for synchrony or to theories in terms of the compensation of differences in neural delays (latencies). This is the main dimension of the Libet discussion. Examples from vision research, such as "temporal-binding-by-synchrony" and the "flash-lag" effect, are then used to illustrate these reasoning patterns in more detail. Alternatively one could assume symbolic representations of time and space (symbolic "tags") that are not coded in their own dimension (not time in time and space in space). Unless such tags are multiplexed with the quality message (tickle, color, or motion), one gets a binding problem for tags. One of the hidden aspects of the discussion between Libet and opponents appears to be the following. Is the NCC smarter than the rest of the nervous system, so that it can solve the problems of local sign (e.g., "where is the event"?) and timing (e.g., "when did it occur?" and "how long did it last?") on its own, or are these pieces of information coded symbolically early on in the system? A supersmart NCC appears to be the assumption of Libet's camp (which includes Descartes, but also mystics). The wish to distribute the smartness evenly across all stages of processing in the nervous system (smart recodings) appears to motivate the opponents. I argue that there are reasons to side with the latter group.     

The disunity of consciousness

Attempts to decode what has become known as the (singular) neural correlate of consciousness (NCC) suppose that consciousness is a single unified entity, a belief that finds expression in the term 'unity of consciousness'. Here, I propose that the quest for the NCC will remain elusive until we acknowledge that consciousness is not a unity, and that there are instead many consciousnesses that are distributed in time and space.     

Temporal binding, binocular rivalry, and consciousness.

Cognitive functions like perception, memory, language, or consciousness are based on highly parallel and distributed information processing by the brain. One of the major unresolved questions is how information can be integrated and how coherent representational states can be established in the distributed neuronal systems subserving these functions. It has been suggested that this so-called "binding problem" may be solved in the temporal domain. The hypothesis is that synchronization of neuronal discharges can serve for the integration of distributed neurons into cell assemblies and that this process may underlie the selection of perceptually and behaviorally relevant information. As we intend to show here, this temporal binding hypothesis has implications for the search of the neural correlate of consciousness. We review experimental results, mainly obtained in the visual system, which support the notion of temporal binding. In particular, we discuss recent experiments on the neural mechanisms of binocular rivalry which suggest that appropriate synchronization among cortical neurons may be one of the necessary conditions for the buildup of perceptual states and awareness of sensory stimuli.     

Information integration based predictions about the conscious states of a spiking neural network

This paper describes how Tononi’s information integration theory of consciousness was used to make detailed predictions about the distribution of phenomenal states in a spiking neural network. This network had approximately 18,000 neurons and 700,000 connections and it used models of emotion and imagination to control the eye movements of a virtual robot and avoid ‘negative’ stimuli. The first stage in the analysis was the development of a formal definition of Tononi’s theory of consciousness. The network was then analysed for information integration and detailed predictions were made about the distribution of consciousness for each time step of recorded activity. This work demonstrates how an artificial system can be analysed for consciousness using a particular theory and in the future this approach could be used to make predictions about the phenomenal states associated with biological systems.     

"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.     

Identification and integration of sensory modalities: Neural basis and relation to consciousness

A key question in studying consciousness is how neural operations in the brain can identify streams of sensory input as belonging to distinct modalities, which contributes to the representation of qualitatively different experiences. The basis for identification of modalities is proposed to be constituted by self-organized comparative operations across a network of unimodal and multimodal sensory areas. However, such network interactions alone cannot answer the question how sensory feature detectors collectively account for an integrated, yet phenomenally differentiated experiential content. This problem turns out to be different from, although related to, the binding problem. It is proposed that the neural correlate of an enriched, multimodal experience is constituted by the attractor state of a dynamic associative network. Within this network, unimodal and multimodal sensory maps continuously interact to influence each other’s attractor state, so that a feature change in one modality results in a fast re-coding of feature information in another modality. In this scheme, feature detection is coded by firing-rate, whereas firing phase codes relational aspects.     

Aristotelian Causation and Neural Correlates of Consciousness

Topoi - Neural correlates of consciousness (NCC) are neural states or processes correlated with consciousness. The aim of this article is to present a coherent explanatory model of NCC that is...     

Conscious behavior explained.

Current neurobiological research on temporal binding in binocular rivalry settings contributes to a better understanding of the neural correlate of perceptual consciousness. This research can easily be integrated into a theory of conscious behavior, but if it is meant to promote a naturalistic theory of perceptual consciousness itself, it is confronted with the notorious explanatory gap argument according to which any statement of psychophysical correlations (and their interpretation) leaves the phenomenal character of, e.g., states of perceptual consciousness open. It is argued that research on temporal binding plays no role in a naturalistic theory of consciousness if the gap argument can be solved on internal philosophical grounds or if it turns out to be unsolvable at the time being. But there may be a way to dissolve or deconstruct it, and the accessibility of this way may well depend on scientific progress, including neurobiological research on the neural correlate of perceptual consciousness.     

The brain is both neurocomputer and quantum computer

In their article, Is the Brain a Quantum Computer,? Litt, Eliasmith, Kroon, Weinstein, and Thagard (2006) criticize the Penrose–Hameroff “Orch OR” quantum computational model of consciousness, arguing instead for neurocomputation as an explanation for mental phenomena. Here I clarify and defend Orch OR, show how Orch OR and neurocomputation are compatible, and question whether neurocomputation alone can physiologically account for coherent gamma synchrony EEG, a candidate for the neural correlate of consciousness. Orch OR is based on quantum computation in microtubules within dendrites in cortex and other regions linked by dendritic–dendritic gap junctions (“dendritic webs”) acting as laterally connected input layers of the brain's neurocomputational architecture. Within dendritic webs, consciousness is proposed to occur as gamma EEG‐synchronized sequences of discrete quantum computational events acting in integration phases of neurocomputational “integrate‐and‐fire” cycles. Orch OR is a viable approach toward understanding how the brain produces consciousness.     

Two theories of consciousness: Semantic pointer competition vs. information integration

Consciousness results from three mechanisms: representation by firing patterns in neural populations, binding of representations into more complex representations called semantic pointers, and competition among semantic pointers to capture the most important aspects of an organism’s current state. We contrast the semantic pointer competition (SPC) theory of consciousness with the hypothesis that consciousness is the capacity of a system to integrate information (IIT). We describe computer simulations to show that SPC surpasses IIT in providing better explanations of key aspects of consciousness: qualitative features, onset and cessation, shifts in experiences, differences in kinds across different organisms, unity and diversity, and storage and retrieval.     

Consciousness, accessibility, and the mesh between psychology and neuroscience

How can we disentangle the neural basis of phenomenal consciousness from the neural machinery of the cognitive access that underlies reports of phenomenal consciousness? We see the problem in stark form if we ask how we can tell whether representations inside a Fodorian module are phenomenally conscious. The methodology would seem straightforward: Find the neural natural kinds that are the basis of phenomenal consciousness in clear cases--when subjects are completely confident and we have no reason to doubt their authority--and look to see whether those neural natural kinds exist within Fodorian modules. But a puzzle arises: Do we include the machinery underlying reportability within the neural natural kinds of the clear cases? If the answer is "Yes," then there can be no phenomenally conscious representations in Fodorian modules. But how can we know if the answer is "Yes"? The suggested methodology requires an answer to the question it was supposed to answer! This target article argues for an abstract solution to the problem and exhibits a source of empirical data that is relevant, data that show that in a certain sense phenomenal consciousness overflows cognitive accessibility. I argue that we can find a neural realizer of this overflow if we assume that the neural basis of phenomenal consciousness does not include the neural basis of cognitive accessibility and that this assumption is justified (other things being equal) by the explanations it allows.     

A Neurofunctional Theory of Visual Consciousness

This paper develops an empirically motivated theory of visual consciousness. It begins by outlining neuropsychological support for Jackendoff's (1987) hypothesis that visual consciousness involves mental representations at an intermediate level of processing. It then supplements that hypothesis with the further requirement that attention, which can come under the direction of high level representations, is also necessary for consciousness. The resulting theory is shown to have a number of philosophical consequences. If correct, higher-order thought accounts, the multiple drafts account, and the widely held belief that sensation precedes perception will all be found wanting. The theory will also be used to illustrate and defend a methodology that fills the gulf between functionalists who ignore the brain and neural reductionists who repudiate functionalism.     

Emotional introspection

One of the most vivid aspects of consciousness is the experience of emotion, yet this topic is given relatively little attention within consciousness studies. Emotions are crucial, for they provide quick and motivating assessments of value, without which action would be misdirected or absent. Emotions also involve linkages between phenomenal and intentional consciousness. This paper examines emotional consciousness from the standpoint of the representational theory of consciousness (RTC). Two interesting developments spring from this. The first is the need for the representation of value, which is distinctive of emotional experience. The second is an extension of RTC's theory of introspection to emotional states, revealing why emotional consciousness is so often introspective even though introspective abilities are not needed to experience emotions, and also explaining why introspection of emotional states is so much less reliable than that of other states of consciousness.     

How could phenomenal consciousness be involved in mental function?

In a phylogenetic perspective, the phenomenal and the functional aspects of consciousness cannot be separated because consciousness, as a phenomenal experience, must be causally effective. The hypothesis I propose is that the fundamental property of consciousness consists of a self-organizing process: the differentiation of a content. The differentiation of a content occurs on the basis of the relations internal to a representational whole, which behaves like a field and tends towards a condition of equilibrium. This hypothesis can be somehow considered an extension of Gestalt visual perceptual theory. Unlike neurocomputational processes, which are non-conscious and extrinsic to the representation, conscious processes are intrinsic to the representational whole. Consciousness, as an intrinsically self-organizing process interwoven with its phenomenal aspects, can be more than epiphenomenal and it can be involved in mental function. The paper then discusses the implications of this hypothesis for subjectivity and the explanatory gap.     

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.