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.     

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.     

Thinking the voice: neural correlates of voice perception

The human voice is the carrier of speech, but also an "auditory face" that conveys important affective and identity information. Little is known about the neural bases of our abilities to perceive such paralinguistic information in voice. Results from recent neuroimaging studies suggest that the different types of vocal information could be processed in partially dissociated functional pathways, and support a neurocognitive model of voice perception largely similar to that proposed for face perception.     

The neural correlates of conceptual and perceptual false recognition

False recognition, broadly defined as a claim to remember something that was not encountered previously, can arise for multiple reasons. For instance, a distinction can be made between conceptual false recognition (i.e., false alarms resulting from semantic or associative similarities between studied and tested items) and perceptual false recognition (i.e., false alarms resulting from physical similarities between studied and tested items). Although false recognition has been associated with frontal cortex activity, it is unclear whether this frontal activity can be modulated by the precise relationship between studied and falsely remembered items. We used event-related fMRI to examine the neural basis of conceptual compared with perceptual false recognition. Results revealed preferential activity in multiple frontal cortex regions during conceptual false recognition, which likely reflected increased semantic processing during conceptual (but not perceptual) memory errors. These results extend recent reports that different types of false recognition can rely on dissociable neural substrates, and they indicate that the frontal activity that is often observed during false compared with true recognition can be modulated by the relationship between studied and tested items.     

Tracking the processes behind conscious perception: a review of event-related potential correlates of visual consciousness

Event-related potential (ERP) studies have attempted to discover the processes that underlie conscious visual perception by contrasting ERPs produced by stimuli that are consciously perceived with those that are not. Variability of the proposed ERP correlates of consciousness is considerable: the earliest proposed ERP correlate of consciousness (P1) coincides with sensory processes and the last one (P3) marks postperceptual processes. A negative difference wave called visual awareness negativity (VAN), typically observed around 200 ms after stimulus onset in occipitotemporal sites, gains strong support for reflecting the processes that correlate with, and possibly enable, aware visual perception. Research suggests that the early parts of conscious processing can proceed independently of top-down attention, although top-down attention may modulate visual processing even before consciousness. Evidence implies that the contents of consciousness are provided by interactions in the ventral stream, but indispensable contributions from dorsal regions influence already low-level visual responses.     

A TMS study of the ventral projections from V1 with implications for the finding of neural correlates of consciousness

The study of subliminal perception in normal and brain lesioned subjects has long been of interest to scholars studying the neural mechanisms behind conscious vision. Using brief durations and a developed methodology of introspective reporting, we present an experiment with visual stimuli that gives rise to little or no subliminal perception under normal viewing conditions. Coupled with transcranial magnetic stimulation, however, we find a dissociation between correctness and conscious awareness. Furthermore, we find support for the hypothesis that the ventral projection streams from V1 are necessary for visual consciousness.     

Structure-from-motion: dissociating perception, neural persistence, and sensory memory of illusory depth and illusory rotation

In the structure-from-motion paradigm, physical motion on a screen produces the vivid illusion of an object rotating in depth. Here, we show how to dissociate illusory depth and illusory rotation in a structure-from-motion stimulus using a rotationally asymmetric shape and reversals of physical motion. Reversals of physical motion create a conflict between the original illusory states and the new physical motion: Either illusory depth remains constant and illusory rotation reverses, or illusory rotation stays the same and illusory depth reverses. When physical motion reverses after the interruption in presentation, we find that illusory rotation tends to remain constant for long blank durations (T _blank ≥ 0.5 s), but illusory depth is stabilized if interruptions are short (T _blank ≤ 0.1 s). The stability of illusory depth over brief interruptions is consistent with the effect of neural persistence. When this is curtailed using a mask, stability of ambiguous vision (for either illusory depth or illusory rotation) is disrupted. We also examined the selectivity of the neural persistence of illusory depth. We found that it relies on a static representation of an interpolated illusory object, since changes to low-level display properties had little detrimental effect. We discuss our findings with respect to other types of history dependence in multistable displays (sensory stabilization memory, neural fatigue, etc.). Our results suggest that when brief interruptions are used during the presentation of multistable displays, switches in perception are likely to rely on the same neural mechanisms as spontaneous switches, rather than switches due to the initial percept choice at the stimulus onset.     

The neural correlates of reward-related trial-and-error learning: an fMRI study with a probabilistic learning task

This fMRI study investigated the neural correlates of reward-related trial-and-error learning in association with changing degrees of stimulus-outcome predictabilities. We found that decreasing predictability was associated with increasing activation in a frontoparietal network. Only maximum predictability was associated with signal decreases across the learning process. The receipt of monetary reward revealed activation in the striatum and associated frontoparietal regions. Present data indicate that during reward-related learning, high uncertainty forces areas relevant for cognitive control to remain activated. In contrast, learning on the basis of predictable stimulus-outcome associations enables the brain to reduce resources in association with the processes of prediction.     

The levels of perceptual processing and the neural correlates of increasing subjective visibility

According to the levels-of-processing hypothesis, transitions from unconscious to conscious perception may depend on stimulus processing level, with more gradual changes for low-level stimuli and more dichotomous changes for high-level stimuli. In an event-related fMRI study we explored this hypothesis using a visual backward masking procedure. Task requirements manipulated level of processing. Participants reported the magnitude of the target digit in the high-level task, its color in the low-level task, and rated subjective visibility of stimuli using the Perceptual Awareness Scale. Intermediate stimulus visibility was reported more frequently in the low-level task, confirming prior behavioral results. Visible targets recruited insulo-fronto-parietal regions in both tasks. Task effects were observed in visual areas, with higher activity in the low-level task across all visibility levels. Thus, the influence of level of processing on conscious perception may be mediated by attentional modulation of activity in regions representing features of consciously experienced stimuli.     

The cognitive and neural correlates of "tactile consciousness": a multisensory perspective

People's awareness of tactile stimuli has been investigated in far less detail than their awareness of stimuli in other sensory modalities. In an attempt to fill this gap, we provide an overview of studies that are pertinent to the topic of tactile consciousness. We discuss the results of research that has investigated phenomena such as "change blindness", phantom limb sensations, and numerosity judgments in tactile perception, together with the results obtained from the study of patients affected by deficits that can adversely affect tactile perception such as neglect, extinction, and numbsense. The similarities as well as some of the important differences that have emerged when visual and tactile conscious information processing have been compared using similar experimental procedures are highlighted. We suggest that conscious information processing in the tactile modality cannot be separated completely from the more general processing of spatial information in the brain. Finally, the importance of considering tactile consciousness within the larger framework of multisensory information processing is also discussed.     

Expanding the discussion: Revision of the fundamental assumptions framing the study of the neural correlates of consciousness

The way one asks a question is shaped by a-priori assumptions and constrains the range of possible answers. We identify and test the assumptions underlying contemporary debates, models, and methodology in the study of the neural correlates of consciousness, which was framed by Crick and Koch’s seminal paper (1990). These premises create a sequential and passive conception of conscious perception: it is considered the product of resolved information processing by unconscious mechanisms, produced by a singular event in time and place representing the moment of entry. The conscious percept produced is then automatically retained to be utilized by post-conscious mechanisms. Major debates in the field, such as concern the moment of entry, the all-or-none vs graded nature, and report vs no-report paradigms, are driven by the consensus on these assumptions. We show how removing these assumptions can resolve some of the debates and challenges and prompt additional questions. The potential non-sequential nature of perception suggests new ways of thinking about consciousness as a dynamic and dispersed process, and in turn about the relationship between conscious and unconscious perception. Moreover, it allows us to present a parsimonious account for conscious perception while addressing more aspects of the phenomenon.     

An fMRI investigation of the neural correlates underlying the processing of novel metaphoric expressions

The neural networks associated with processing related pairs of words forming literal, novel, and conventional metaphorical expressions and unrelated pairs of words were studied in a group of 15 normal adults using fMRI. Subjects read the four types of linguistic expressions and decided which relation exists between the two words (metaphoric, literal, or unrelated). According to the Graded Salience Hypothesis (GSH, ), which predicts a selective RH involvement in the processing of novel, nonsalient meanings, it is primarily the degree of meaning salience of a linguistic expression rather than literality or nonliterality, which modulates the degree of left hemisphere (LH) and right hemisphere (RH) processing of metaphors. In the present study, novel metaphorical expressions represented the nonsalient interpretations, whereas conventional metaphors and literal expressions represented the salient interpretations. A direct comparison of the novel metaphors vs. the conventional metaphors revealed significantly stronger activity in right posterior superior temporal sulcus, right inferior frontal gyrus, and left middle frontal gyrus. These results support the GSH and suggest a special role for the RH in processing novel metaphors. Furthermore, the right PSTS may be selectively involved in verbal creativity.     

The neural correlates of statistical learning in a word segmentation task: An fMRI study

Functional magnetic resonance imaging (fMRI) was used to assess neural activation as participants learned to segment continuous streams of speech containing syllable sequences varying in their transitional probabilities. Speech streams were presented in four runs, each followed by a behavioral test to measure the extent of learning over time. Behavioral performance indicated that participants could discriminate statistically coherent sequences (words) from less coherent sequences (partwords). Individual rates of learning, defined as the difference in ratings for words and partwords, were used as predictors of neural activation to ask which brain areas showed activity associated with these measures. Results showed significant activity in the pars opercularis and pars triangularis regions of the left inferior frontal gyrus (LIFG). The relationship between these findings and prior work on the neural basis of statistical learning is discussed, and parallels to the frontal/subcortical network involved in other forms of implicit sequence learning are considered.     

On the neural correlates of object recognition awareness: relationship to computational activities and activities mediating perceptual awareness

Based on theoretical considerations of Aurell (1979) and Block (1995), we argue that object recognition awareness is distinct from purely sensory awareness and that the former is mediated by neuronal activities in areas that are separate and distinct from cortical sensory areas. We propose that two of the principal functions of neuronal activities in sensory cortex, which are to provide sensory awareness and to effect the computations that are necessary for object recognition, are dissociated. We provide examples of how this dissociation might be achieved and argue that the components of the neuronal activities which carry the computations do not directly enter the awareness of the subject. The results of these computations are sparse representations (i.e., vector or distributed codes) which are activated by the presentation of particular sensory objects and are essentially engrams for the recognition of objects. These final representations occur in the highest order areas of sensory cortex; in the visual analyzer, the areas include the anterior part of the inferior temporal cortex and the perirhinal cortex. We propose, based on lesion and connectional data, that the two areas in which activities provide recognition awareness are the temporopolar cortex and the medial orbitofrontal cortex. Activities in the temporopolar cortex provide the recognition awareness of objects learned in the remote past (consolidated object recognition), and those in the medial orbitofrontal cortex provide the recognition awareness of objects learned in the recent past. The activation of the sparse representation for a particular sensory object in turn activates neurons in one or both of these regions of cortex, and it is the activities of these neurons that provide the awareness of recognition of the object in question. The neural circuitry involved in the activation of these representations is discussed.     

Visual marking: dissociating effects of new and old set size

Visual marking makes it possible to ignore old items during search. In a typical study, old items are previewed 1 s before adding an equal number of new items, one of which is the target. Previewing half of the items reduces the search slope relating response time (RT) to overall set size by half. However, this manipulation sometimes only reduces overall RT but not search slope (Experiment 1). By orthogonally varying the numbers of old and new items, Experiment 2 shows that old and new set sizes interactively affect visual marking. Given a constant new set size, the size of the old set has negligible effect on RT. However, increasing the new set size reduces the preview benefit in overall RT. Experiment 3 shows that this reduction may be restricted to paradigms that use temporal segregation cues. Studies should vary old and new set size orthogonally to avoid missing a visual marking effect where one may be present.     

The neural correlates of happiness: A review of PET and fMRI studies using autobiographical recall methods

Although very difficult to define, happiness is becoming a core concept within contemporary psychology and affective neuroscience. In the last two decades, the increased use of neuroimaging techniques has facilitated empirical study of the neural correlates of happiness. This area of research utilizes procedures that induce positive emotion and mood, and autobiographical recall is one of the most widely used and effective approaches. In this article, we review eight positron emission tomography and seven functional magnetic resonance imaging studies that have investigated happiness by using autobiographical recall to induce emotion. Regardless of the neuroimaging technique used, the studies conducted so far have shown that remembering happy events is primarily associated with the activation of many areas, including anterior cingulate cortex, prefrontal cortex, and insula. Importantly, these areas are also found to be connected with other basic emotions, such as sadness and anger. In the conclusion, we integrate these findings, discussing important limitations of the extant literature and suggesting new research directions.