How to Find the Neural Correlate of Consciousness

Afterimages: A Tool for Defining the Neural Correlate of Visual Consciousness

Identifying neural correlates of visual consciousness with ALE meta-analyses

OPINION article Front. Hum. Neurosci., 19 July 2013Sec. Cognitive Neuroscience Volume 7 - 2013 | https://doi.org/10.3389/fnhum.2013.00387

EDITORIAL article Front. Psychol., 01 September 2011 | https://doi.org/10.3389/fpsyg.2011.00191

Subjective experience has often taken center stage in debates between competing conceptual theories of the mind. This is also a central object of concern in the empirical domain, and especially in the search for the neural correlates of consciousness (NCCs). By now, most of the competing conceptual theories of consciousness have become aligned with distinct hypotheses about the NCCs. These hypotheses are usually distinguished by reference to their proposed location of the NCCs. This difference in hypothesized location of the NCCs has tempted participants in these debates to infer that evidence indicating the location of the NCCs in one or the other brain region can be taken as direct evidence for or against a given conceptual theory of consciousness. We argue that this is an overestimation of the work finding the NCCs can do for us, and that there are principled reasons to resist this kind of inference. To show this we point out the lack of both an isomorphism and a homomorphism between the conceptual frameworks in which most theories are cached, and the kind of data we can get from neuroimaging. The upshot is that neural activation profiles are insufficient to distinguish between competing theories in the conceptual domain. We suggest two ways to go about ameliorating this issue.

An increasing number of authors suggest that the neural correlates of consciousness (NCC) have no selective, executive, or metacognitive function. It is believed that attention unconsciously selects the contents that will become conscious. Consciousness would have only the fundamental function of transforming the selected contents into a format easily used by high-level processors, such as working memory, language, or autobiographical memory. According to Dehaene, the neural correlates (NC) of access consciousness (AC; cognitive consciousness) constitute a widespread network in the frontal, parietal, and temporal cortices. While Tononi localized the correlates of phenomenal consciousness (PC; subjective consciousness) to a posterior “hot zone” in the temporo-parietal cortex. A careful examination of the works of these two groups leads to the conclusion that the correlates of access and PC coincide. The two consciousnesses are therefore two faces of the same single consciousness with both its cognitive and subjective contents. A review of the literature of the pathology called “neglect” confirms that the common correlates include 10: a memory center, an activation center, and eight parallel centers. From study of the “imagery” it can be deduced that these eight parallel centers would operate as points of convergence in the third person linking the respective eight sensory-motor-emotional areas activated by external perceptions and the corresponding memories of these perceptions deposited in the memory center. The first four centers of convergence appear in the most evolved fish and gradually reach eight in humans.

Experimental and theoretical approaches aiming at the establishment of neural correlates of higher cognitive functions and awareness have been extensively studied in the past decade. Information-theoretical indices are useful tools in establishing quantitative metrics when analyzing data of cognitive experiments. In this work we report a systematic statistical analysis of multiple runs of ECoG measurements over the rabbit visual cortex. The results are interpreted invoking the concept of Pragmatic Information, which is complementary to the Shannon Entropy Index. We interpret these finding based on a dynamical system approach to brains and cognition. We identify large-scale synchronization across broad frequency band as potential manifestation of the `aha' effect, indicating the construction of knowledge and meaning from input sensory data and leading to awareness experience.

For many years, since Baars (1988) explicitly formulated it, contrastive analysis has been the key methodological approach in experimental studies of consciousness. When certain properly chosen psychological experimental setups (allowing an invariant target stimulus either to be consciusly experienced or not) were combined with brain-imaging methods, contrastive analysis became a quite powerful tool of research (Crick, 1994; Koch, 2004). By subtracting markers of brain processes recorded in the conditions without conscious experience of the target from the markers recorded in the conditions where the same target is consciously experienced it was believed that the markers of neural correlates of consciousness (NCC) can be obtained. However, as it turned out in the subsequent theoretical and experimental analysis, the picture is not so clear and simple (Bachmann, 2000, 2009; Miller, 2007; Aru et al., 2012; de Graaf et al., 2012). For example, when in the invariant conditions of independent variables a masked visual stimulus was consciously perceived or not (consciousness of the target standed as a dependent variable), NCC which were measured as a spectral perturbation of EEG was present already before stimulus presentation (Aru and Bachmann, 2009). Thus, the neural correlate of consciousness of a stimulus was present earlier than the stimulus itself was presented. Now, a reader must not get excited here because instead of some paranormal explanations brain-science based explanations can be comfortably used. In order to overcome the conceptual crisis hitting the traditional contrastive analysis based NCC research it was suggested that unconscious prerequisite processes (NCCpr) emerging as a result of contrastive analysis of brain-process markers of consciousness and similarly unconscious consequent processes (NCCco) must be differentiated from the constitutive processes directly associated with conscious experience (Aru et al., 2012; de Graaf et al., 2012). Thus, new experimental approaches were in need to avoid the trap of distilling prerequisite, direct, and consequent processes as mutually confounded and empirically inseparable. Despite some first attempts in this direction (Aru and Bachmann, 2015), the specialist landscape in this domain has remained obscure and no breakthrough solutions have been in sight. Moreover, there seems to be a number of additional uncertainties when we try to disentangle the various sub-types of NCC. Even NCCpr and NCCco are not unitary in terms of their theoretical meaning and associated neural processes. First, as the contents on which the perceptual report is founded can be selective, the markers of unused conscious contents may be erroneously neglected as markers of unconscious processes. They actually belong to consciousness level processes, but related to contents of consciousness qualitatively different from the ones specified by NCC. Second, in measuring NCC we must be able to disentangle contributions of the general consciousness enabling mechanisms and the selective contents representing mechanisms because their markers can be different and thus confused. In what follows I will substantiate these two issues.

Two neural correlates of consciousness

A global workspace model for phenomenal and access consciousness

'Self' and the Problem of Consciousness

This article investigates neural and physiological correlates of simulator sickness (SS) through a controlled experiment conducted within a fully immersive dome projection system. Our goal is to establish a reliable, objective, and in situ measurable predictive indicator of SS. SS is a problem common to all types of visual simulators consisting of motion sickness‐like symptoms that may be experienced while and after being exposed to a dynamic, immersive visualization. It leads to ethical concerns and impaired validity of simulator‐based research. Due to the popularity of virtual reality devices, the number of people exposed to this problem is increasing and, therefore, it is crucial to find reliable predictors of this condition before any symptoms appear. Despite its relevance and the several theories about its origins, SS cannot yet be quantitatively modeled and predicted. Our results indicate that, while neural correlates did not materialize, physiological measures may be a solid early indicator of oncoming SS.

1. With the use of two independent microelectrodes and multiunit spike separation, 26 neural pairs, 17 triplets, and 8 quadruplets were recorded in the auditory midbrain of the leopard frog, resulting in a total of 125 neural pairs. 2. Functional interrelationships between neurons were studied by analyzing 638 cross-coincidence histograms as functions of stimulus type, stimulus level, and estimated neuron distance. Significance criteria for correlograms were established on the basis of the distribution of extreme values in a large number of correlograms for nonsimultaneously recorded pairs. 3. Simultaneous recordings from three neurons, that all showed significant neural pair correlations were analyzed with the use of the joint occurrence diagram, which displays the joint coincidences for the firings of two units (a and b) with the firings of the trigger unit (c). 4. It was found that 97.5% of the pairs showed a significant stimulus-induced correlation; neighboring neurons exhibited a stronger stimulus correlation (synchrony) than more distant neurons. 5. Positive neural interaction strength (75% to shared excitatory input) was independent of neuron distance (taking into account that the estimated electrode distance in the present investigation was never greater than 300 microns) and occurred in 25% of the pairs investigated. About 25% of the positive neural correlations could be attributed to unidirectional excitation, the majority of which was found for single-electrode pairs. Negative neural correlation occurred in 8% of the pairs and, with one exception, was found only for neurons recorded on the same electrode. 6. Evidence for the presence of feed-forward and/or feedback inhibition was found. 7. There was a strong stimulus-type influence on stimulus correlation and on positive neural correlation, whereas stimulus intensity affected the stimulus correlation but not the neural correlation. 8. From the incidence of triplet correlations, it was concluded that the divergence of afferents onto midbrain neurons was limited; it was unlikely that more than three neurons were contacted by one afferent. In contrast, convergence of afferents on torus semicircularis cells was widespread; 40-50% of the midbrain neurons were bimodally tuned and received input originating from the two auditory papillae. Convergence of fibers from the same papilla was also extensive. 9. Fast modulation of functional neural connectivity through the activity of other neurons was found, although this was probably not the result of actual changes in synaptic strength but of synchronized changes in firing rate.

To investigate neural and behavioral correlates of emotional experiences as potential vulnerability markers in remitted depression. Fourteen remitted participants with a history of major depression and fourteen closely matched healthy control participants took part in the study. We used two psychiatric interviews (Hamilton Depression Rating Scale, Montgomery-Asberg Depression Rating Scale) and one self-report scale (Beck Depression Inventory) to assess remission. Healthy control participants were interviewed by an experienced psychiatrist to exclude those who showed any current or lifetime psychiatric or neurological disorders. To explore psychosocial and cognitive-interpersonal underpinnings of potential vulnerability markers of depression, early life stress, coping styles and alexithymia were also assessed. We induced pleasant and unpleasant emotional states using congruent combinations of music and human emotional faces to investigate neural and behavioral correlates of emotional experiences; neutral stimuli were used as a control condition. Brain responses were recorded using functional magnetic resonance imaging. Behavioral responses of pleasantness, arousal, joy and fear were measured via button-press inside the resonance imaging scanner. The mean age of the sample was 54.9 (± 11.3) years. There were no differences between remitted depressed (RD) (n = 14; 9 females and 5 males) and healthy participants (n = 14; 8 females and 6 males) regarding age, current degree of depression, early life stress, coping styles and alexithymia. On a neural level, RD participants showed reduced activations in the pregenual anterior cingulate cortex (pgACC) in response to pleasant [parameter estimates: -0.78 vs 0.32; t(26) = -3.41, P < 0.05] and unpleasant [parameter estimates: -0.88 vs 0.56; t(26)= -4.02, P < 0.05] emotional stimuli. Linear regression analysis revealed that pgACC activity was modulated by early life stress [β = -0.48; R(2) = 0.23, F(1,27) = 7.83, P < 0.01] and task-oriented coping style [β = 0.63; R(2) = 0.37, F(1,27) = 16.91, P < 0.001]. Trait anxiety modulated hippocampal responses to unpleasant stimuli [β = 0.62; R(2) = 0.38, F(1,27) = 15.95, P < 0.001]. Interestingly, in their reported experiences of pleasantness, arousal, happiness and fear in response to pleasant, unpleasant and neutral stimuli, RD participants did not differ significantly from healthy control participants. Adding trait anxiety or alexithymia as a covariate did not change the results. The present study indicates that, in euthymic individuals, depression history alters neural correlates, but not the subjective dimension of pleasant and unpleasant emotional experiences.

Twenty years ago, Thomas Metzinger published the book "The Neural Correlates of Consciousness" amassing the state of knowledge in the field of consciousness studies at the time from philosophical and empirical perspectives. On the occasion of the 20th anniversary of this impactful publication, we review the progress the field has made since then and the important methodological challenges it faces. A tremendous number of empirical studies have been conducted, which has led to the identification of many candidate neural correlates of consciousness. Yet, this tremendous amount of work has not unraveled a consensual account of consciousness as of now. Many questions, some already raised twenty years ago, remain unanswered, and an enormous proliferation of theories sharply contrasts with the scarcity of compelling data and methodological challenges. The contrastive method, the foundational method used to study the neural correlate of consciousness (NCC), has also been called into question. And while awareness in the community of its shortcomings is widespread, few concrete attempts have been made to go beyond it and/or to revise existing theories. We propose several methodological shifts that we believe may help to advance the quest of the NCC program, while remaining uncommitted to any specific theory: (1) the currently prevalent “contrastive method” should lose its monopoly in favor of methods that attempt to explain the phenomenology of experience; (2) experimental data should be shared in centralized, multi-methods databases, transcending the limitations of individual experiments by granting granularity and power to generalize findings and “distill” the NCC proper; (3) the explanatory power of theories should be directly pitted against each other to overcome the non-productive fractioning of the field into insular camps seeking confirmatory evidence for their theories. We predict these innovations might enable the field to progress towards the goal of explaining consciousness.