We present a theoretical view of the cellular foundations for network-level processes involved in producing our conscious experience. Inputs to apical synapses in layer 1 of a large subset of neocortical cells are summed at an integration zone near the top of their apical trunk. These inputs come from diverse sources and provide a context within which the transmission of information abstracted from sensory input to their basal and perisomatic synapses can be amplified when relevant. We argue that apical amplification (...) enables conscious perceptual experience and makes it more flexible, and thus more adaptive, by being sensitive to context. Apical amplification provides a possible mechanism for recurrent processing theory that avoids strong loops. It makes the broadcasting hypothesized by global neuronal workspace theories feasible while preserving the distinct contributions of the individual cells receiving the broadcast. It also provides mechanisms that contribute to the holistic aspects of integrated information theory. As apical amplification is highly dependent on cholinergic, aminergic, and other neuromodulators, it relates the specific contents of conscious experience to global mental states and to fluctuations in arousal when awake. We conclude that apical dendrites provide a cellular mechanism for the context-sensitive selective amplification that is a cardinal prerequisite of conscious perception. (shrink)

A central debate in philosophy and neuroscience pertains to whether PFC activity plays an essential role in the neural basis of consciousness. Neuroimaging and electrophysiology studies have revealed that the contents of conscious perceptual experience can be successfully decoded from PFC activity, but these findings might be confounded by post- perceptual cognitive processes, such as thinking, reasoning, and decision-making, that are not necessary for con- sciousness. To clarify the involvement of the PFC in consciousness, we present a synthesis of research (...) that has used intracranial electrical stimulation (iES) for the causal modulation of neural activity in the human PFC. This research provides compelling evidence that iES of only certain prefrontal regions (i.e., orbitofrontal cortex and anterior cingu- late cortex) reliably perturbs conscious experience. Conversely, stimulation of anterolateral prefrontal sites, often con- sidered crucial in higher-order and global workspace theories of consciousness, seldom elicits any reportable alterations in consciousness. Furthermore, the wide variety of iES-elicited effects in the PFC (e.g., emotions, thoughts, and olfactory and visual hallucinations) exhibits no clear relation to the immediate environment. Therefore, there is no evidence for the kinds of alterations in ongoing perceptual experience that would be predicted by higher-order or global workspace theories. Nevertheless, effects in the orbitofrontal and anterior cingulate cortices suggest a specific role for these PFC subregions in supporting emotional aspects of conscious experience. Overall, this evidence presents a challenge for higher-order and global workspace theories, which commonly point to the PFC as the basis for con- scious perception based on correlative and possibly confounded information. (shrink)

Working memory, an important posit in cognitive science, allows one to temporarily store and manipulate information in the service of ongoing tasks. Working memory has been traditionally classified as an explicit memory system – that is, as operating on and maintaining only consciously perceived information. Recently, however, several studies have questioned this assumption, purporting to provide evidence for unconscious working memory. In this paper, we focus on visual working memory and critically examine these studies as well as studies of unconscious (...) perception that seem to provide indirect evidence for unconscious working memory. Our analysis indicates that current evidence does not support an unconscious working memory store, though we offer independent reasons to think that working memory may operate on unconsciously perceived information. (shrink)

Consciousness is scientifically challenging to study because of its subjective aspect. This leads researchers to rely on report-based experimental paradigms in order to discover neural correlates of consciousness (NCCs). I argue that the reliance on reports has biased the search for NCCs, thus creating what I call 'methodological artefacts'. This paper has three main goals: first, describe the measurement problem in consciousness science and argue that this problem led to the emergence of methodological artefacts. Second, provide a critical assessment of (...) the NCCs put forward by the global neuronal workspace theory. Third, provide the means of dissociating genuine NCCs from methodological artefacts. (shrink)

Research into the neural correlates of consciousness (NCCs) has blossomed, due to the advent of new and increasingly sophisticated brain research tools. Neuroimaging has uncovered a variety of brain processes that relate to conscious perception, obtained in a range of experimental paradigms. But methods such as functional magnetic resonance imaging or electroencephalography do not always afford inference on the functional role these brain processes play in conscious vision. Such empirical NCCs could reflect neural prerequisites, neural consequences, or neural substrates of (...) a conscious experience. Here, we take a closer look at the use of non-invasive brain stimulation (NIBS) techniques in this context. We discuss and review how NIBS methodology can enlighten our understanding of brain mechanisms underlying conscious vision by disentangling the empirical NCCs. (shrink)

The interocular suppression technique termed continuous flash suppression (CFS) has become an immensely popular tool for investigating visual processing outside of awareness. The emerging picture from studies using CFS is that extensive processing of a visual stimulus, including its semantic and affective content, occurs despite suppression from awareness of that stimulus by CFS. However, the current implementation of CFS in many studies examining processing outside of awareness has several drawbacks that may be improved upon for future studies using CFS. In (...) this paper, we address some of those shortcomings, particularly ones that affect the assessment of unawareness during CFS, and ones to do with the use of ‘visible’ conditions that are often included as a comparison to a CFS condition. We also discuss potential biases in stimulus processing as a result of spatial attention and feature-selective suppression. We suggest practical guidelines that minimize the effects of those limitations in using CFS to study visual processing outside of awareness. (shrink)

This review addresses the role of early sensory areas in the awareness of tactile information in humans. The results of recent studies dealing with this important topic are critically discussed: In particular, we report on evidence from neuropsychology, neurophysiology, neuroimaging, and behavioral experiments that have highlighted the crucial role played by the primary somatosensory cortex in mediating our awareness of tactile information. Phenomena, such as tactile hallucinations, tactile illusions, the perception of supernumerary limbs, and synaesthesia are also discussed. The research (...) reviewed here clearly shows that the activation of SI is necessary, but not sufficient, for the awareness of touch. On the basis of the evidence outlined here, we propose a neurocognitive model that provides a conceptual framework in which to interpret the results of the literature regarding tactile consciousness. (shrink)

Two types of the attentional network, alerting and orienting, help organisms respond to environmental events for survival in the temporal and spatial dimensions, respectively. Here, we applied chromatic flicker beyond the critical fusion frequency to address whether awareness was necessary for activation of the two attentional networks. We found that high-frequency chromatic flicker, despite its failure to reach awareness, produced the alerting and orienting effects, supporting the dissociation between attention and awareness. Furthermore, as the flicker frequency increased, the orienting effect (...) attenuated whereas the alerting effect remained unchanged. According to the systematic decline in temporal frequency sensitivity across the visual hierarchy, this finding suggests that unconscious alerting might be associated with activity in earlier visual areas than unconscious orienting. Since high-frequency flicker has been demonstrated to only activate early visual cortex, we suppose that neural activation in early visual areas might be sufficient to activate the two attentional networks. (shrink)

People can maintain accurate representations of visual changes without necessarily being aware of them. Here, we investigate whether a similar phenomenon also exists in touch. In Experiments 1 and 2, participants detected the presence of a change between two consecutively-presented tactile displays. Tactile change blindness was observed, with participants failing to report the presence of tactile change. Critically, however, when participants had to make a forced choice response regarding the number of stimuli presented in the two displays, their performance was (...) significantly better than chance . Experiment 3 demonstrated that tactile change detection does not necessarily involve a shift of spatial attention toward the location of change, regardless of whether the change is explicitly detected. We conclude that tactile change detection likely results from comparing representations of the two displays, rather than by directing spatial attention to the location of the change. (shrink)

Conscious visual perception can fail in many circumstances. However, little is known about the causes and processes leading to failures of visual awareness. In this study, we introduce a new signal detection measure termed subjective discriminability of invisibility that allows one to distinguish between subjective blindness due to reduction of sensory signals or to lack of attentional access to sensory signals. The SDI is computed based upon subjective confidence in reporting the absence of a target . Using this new measure, (...) we found that target misses were subjectively indistinguishable from physical absence when contrast reduction, backward masking and flash suppression were used, whereas confidence was appropriately modulated when dual task, attentional blink and spatial uncertainty methods were employed. These results show that failure of visual perception can be identified as either a result of perceptual or attentional blindness depending on the circumstances under which visual awareness was impaired. (shrink)

Previous research has shown implicit semantic processing of faces or pictures, but whether symbolic carriers such as words can be processed this way remains controversial. Here we examine this issue by adopting the continuous flash suppression paradigm to ensure that the processing undergone is indeed unconscious without the involvement of partial awareness. Negative or neutral words projected into one eye were made invisible due to strong suppression induced by dynamic-noise patterns shown in the other eye through binocular rivalry. Inverted and (...) scrambled words were used as controls to provide baselines at orthographic and feature levels, respectively. Compared to neutral words, emotion-described and emotion-induced negative words required longer time to release from suppression, but only for upright words. These results suggest that words can be processed unconsciously up to semantic level since under interocular suppression completely invisible words can lead to different processing speed due to the emotion information they carry. (shrink)

In the cognitive neuroscience of consciousness, participants have commonly been instructed to report their conscious content. This, it was claimed, risks confounding the neural correlates of consciousness (NCC) with their preconditions, i.e., allocation of attention, and consequences, i.e., metacognitive reflection. Recently, the field has therefore been shifting towards no-report paradigms. No-report paradigms draw their validity from a direct comparison with no-report conditions. We analyze several examples of such comparisons and identify alternative interpretations of their results and/or methodological issues in all (...) cases. These go beyond the previous criticism that just removing the report is insufficient, because it does not prevent metacognitive reflection. The conscious mind is fickle. Without having much to do, it will turn inward and switch, or timeshare, between the stimuli on display and daydreaming or mind-wandering. Thus, rather than the NCC, no-report paradigms might be addressing the neural correlates of conscious disengagement. This observation reaffirms the conclusion that no-report paradigms are no less problematic than report paradigms. (shrink)

Bistable figures provide a fascinating window through which to explore human visual awareness. Here we demonstrate for the first time that the semantic context provided by a background auditory soundtrack can modulate an observer’s predominant percept while watching the bistable “my wife or my mother-in-law” figure . The possibility of a response-bias account—that participants simply reported the percept that happened to be congruent with the soundtrack that they were listening to—was excluded in Experiment 2. We further demonstrate that this crossmodal (...) semantic effect was additive with the manipulation of participants’ visual fixation , while it interacted with participants’ voluntary attention . These results indicate that audiovisual semantic congruency constrains the visual processing that gives rise to the conscious perception of bistable visual figures. Crossmodal semantic context therefore provides an important mechanism contributing to the emergence of visual awareness. (shrink)

People with grapheme-color synesthesia perceive specific colors when viewing different letters or numbers. Previous studies have suggested that synesthetic color experience can be bistable when induced by an ambiguous character. However, the exact relationship between processes underlying the identity of an alphanumeric character and the experience of the induced synesthetic color has not been examined. In the present study, we explored this by focusing on the temporal relation of inducer identification and color emergence using inducers whose identity could be rendered (...) ambiguous upon rotation of the characters. Specifically, achromatic alphabetic letters and digits were presented at varying angles to 9 grapheme-color synesthetes. Results showed that grapheme identification and synesthetically perceived grapheme color covary with the orientation of the test stimulus and that synesthetes were slower naming the experienced color than identifying the character, particularly at intermediate angles where ambiguity was greatest. (shrink)

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. (shrink)

Strawson’s challenging and provocative defence of panpsychism1 begins by sensibly insisting that physicalism, properly understood, must unflinchingly countenance the occurrence of conscious experiences. No view, he urges, will count as ‘real physicalism’ (p. 4) if it seeks to get around or soften that commitment, as versions of socalled physicalism sometimes do. Real physicalism (hereinafter physicalism tout court) must accordingly reject any stark opposition of mental and physical, which is not only invoked by many followers of Descartes, but even countenanced by (...) many recent physicalists. Conscious experiences, Strawson persuasively urges, are a special case of the physical, just as cows are animals. Panpsychism enters the picture because, despite the physical nature of conscious experiences, Strawson maintains that we cannot describe or explain the experiential using the terms of physics or neurophysiology. Since the experiential is nonetheless physical, Strawson concludes that the physical ultimates must, in addition to whatever properties physics and neurophysiology reveal, have experiential properties as well. Strawson argues that we cannot avoid this conclusion by maintaining that combinations of physical ultimates constitute or give rise to experiential properties. Physical ultimates cannot give rise to conscious experience unless those ultimates are themselves in some way ‘intrinsically experiential’ (p. 22). The experiential cannot emerge from nonexperiential ultimates in the way that macroscopic liquidity is standardly held to emerge from molecular properties. Strawson concludes that at least some physical ultimates must be ‘intrinsically experience-involving’ (p. 22); and, since it’s reasonable to see the physical ultimates as homogeneous in nature, we should assume that they all have experiential properties. (shrink)

Do perceptual experiences always inherit the content of their neural correlates? Most scientists and philosophers working on perception say ‘yes’. They hold the view that an experience’s content just is (i.e. is identical to) the content of its neural correlate. This paper presses back against this view, while trying to retain as much of its spirit as possible. The paper argues that type-2 blindsight experiences are plausible cases of experiences which lack the content of their neural correlates. They are not (...) experiences of the stimuli or stimulus properties prompting them, but their neural correlates represent these stimulus properties. The argument doesn’t depend on any special view of what it is for an experience to be of a stimulus or stimulus property. The upshot is that, even assuming there is a deep relationship between experiential content and neural content, that relationship is more complex than simple identity. (shrink)

Recently, the mechanistic framework of active inference has been put forward as a principled foundation to develop an overarching theory of consciousness which would help address conceptual disparities in the field (Wiese 2018 ; Hohwy and Seth 2020 ). For that promise to bear out, we argue that current proposals resting on the active inference scheme need refinement to become a process theory of consciousness. One way of improving a theory in mechanistic terms is to use formalisms such as computational (...) models that implement, attune and validate the conceptual notions put forward. Here, we examine how computational modelling approaches have been used to refine the theoretical proposals linking active inference and consciousness, with a focus on the extent and success to which they have been developed to accommodate different facets of consciousness and experimental paradigms, as well as how simulations and empirical data have been used to test and improve these computational models. While current attempts using this approach have shown promising results, we argue they remain preliminary in nature. To refine their predictive and structural validity, testing those models against empirical data is needed i.e., new and unobserved neural data. A remaining challenge for active inference to become a theory of consciousness is to generalize the model to accommodate the broad range of consciousness explananda; and in particular to account for the phenomenological aspects of experience. Notwithstanding these gaps, this approach has proven to be a valuable avenue for theory advancement and holds great potential for future research. (shrink)

Neurophysiological studies in humans employing magneto- and electro- encephalography increasingly suggest that oscillatory rhythmic activity of the brain may be a core mechanism for binding sensory information across space, time, and object features to generate a unified perceptual representation. To distinguish whether oscillatory activity is causally related to binding processes or whether, on the contrary, it is a mere epiphenomenon, one possibility is to employ neuromodulatory techniques such as transcranial alternating current stimulation. tACS has seen a rising interest due to (...) its ability to modulate brain oscillations in a frequency-dependent manner. In the present review, we critically summarize current tACS evidence for a causal role of oscillatory activity in spatial, temporal, and feature binding in the context of visual perception. For temporal binding, the emerging picture supports a causal link with the power and the frequency of occipital alpha rhythms ; however, there is no consistent evidence on the causal role of the phase of occipital tACS. For feature binding, the only study available showed a modulation by occipital alpha tACS. The majority of studies that successfully modulated oscillatory activity and behavioral performance in spatial binding targeted parietal areas, with the main rhythms causally linked being the theta and beta frequency bands. On the other hand, spatio-temporal binding has been directly modulated by parieto-occipital gamma and alpha tACS, suggesting a potential role of cross-frequency coupling when binding across space and time. Nonetheless, negative or partial results have also been observed, suggesting methodological limitations that should be addressed in future research. Overall, the emerging picture seems to support a causal role of brain oscillations in binding processes and, consequently, a certain degree of plasticity for shaping binding mechanisms in visual perception, which, if proved to have long lasting effects, can find applications in different clinical populations. (shrink)

Contour adaptation is a recently described paradigm that renders otherwise salient visual stimuli temporarily perceptually invisible. Here we investigate whether this illusion can be exploited to study visual awareness. We found that CA can induce seconds of sustained invisibility following similarly long periods of uninterrupted adaptation. Furthermore, even fragmented adaptors are capable of producing CA, with the strength of CA increasing monotonically as the adaptors encompass a greater fraction of the stimulus outline. However, different types of adaptor patterns, such as (...) distinctive shapes or illusory contours, produce equivalent levels of CA suggesting that the main determinants of CA are low-level stimulus characteristics, with minimal modulation by higher-order visual processes. Taken together, our results indicate that CA has desirable properties for studying visual awareness, including the production of prolonged periods of perceptual dissociation from stimulation as well as parametric dependencies of that dissociation on a host of stimulus parameters. (shrink)

In general, stimuli that are familiar and recognizable have an advantage of predominance during binocular rivalry. Recent research has demonstrated that familiar and recognizable stimuli such as upright faces and words in a native language could break interocular suppression faster than their matched controls. In this study, a visible word prime was presented binocularly then replaced by a high-contrast dynamic noise pattern presented to one eye and either a semantically related or unrelated word was introduced to the other eye. We (...) measured how long it took for target words to break from suppression. To investigate word-parts priming, a second experiment also included word pairs that had overlapping subword fragments. Results from both experiments consistently show that semantically related words and words that shared subword fragments were faster to gain dominance compared to unrelated words, suggesting that words, even when interocularly suppressed and invisible, can benefit from semantic and subword priming. (shrink)

Consciousness researchers are interested in distinguishing between mental activity that occurs with and without awareness . The inattentional blindness paradigm is an excellent tool for this question because it permits the independent manipulation of processing time and awareness. In the present study, we show that implicit texture segregation can occur during inattentional blindness, provided that the texture is exposed for a sufficient duration. In contrast, a Simon effect does not occur during inattentional blindness, even with similar exposure duration of the (...) critical stimulus. This reveals a dissociation between processing time and awareness, consistent with the possibility of separate processing streams for explicit and implicit visual perception. (shrink)

Mental causation is a predominantly theoretical topic rather than a topic studied in the laboratory. The purpose of this paper is to outline a general approach for studying mental causation by empirical means for philosophers and scientists interested in the topic. The aim is to outline how we can infer mental causation by empirical methods given an unknown solution to the mind-body problem. The approach is based on the principles of causal inference to find causal relations among observed variables used (...) in all branches of science. With these principles, it is possible to estimate the causal effects of mental events: Make an experimental manipulation of a mental event, control confounding variables, and estimate causal effects on the outcome. The caveat is that we cannot separate the causal effects of a mental event from the physical base of the mental event, independent of whether we assume mental events can be reduced to their physical base. A challenge to estimating causal effects of mental events is that measured physical variables, such as electrophysiological potentials from the brain, might reflect processes that are part of “higher-order” phenomena, such as mental events. This means that controlling “downwards” for confounding variables is challenging. It is, however, not impossible. It also means that inferring non-mental causes of action cannot be done by measuring only physical variables alone. Keeping the mind-body problem in mind when designing experiments, it is possible to infer mental causation. (shrink)