Neural signs and mechanisms of consciousness: Is there a potential convergence of theories of consciousness in sight?

Various theories for the neural basis of consciousness have been proposed, suggesting a diversity of neural signs and mechanisms. We ask to what extent this diversity is real, or whether many theories share the same basic ideas with a potential for convergence towards a more unified theory of the neural basis of consciousness. For that purpose, we review and compare the various neural signs, measures, and mechanisms proposed in the different theories.

Split-Brain: What We Know Now and Why This is Important for Understanding Consciousness.

Recently, the discussion regarding the consequences of cutting the corpus callosum ("split-brain") has regained momentum (Corballis, Corballis, Berlucchi, & Marzi, Brain, 141(6), e46, 2018; Pinto et al., Brain, 140(5), 1231-1237, 2017a; Pinto, Lamme, & de Haan, Brain, 140(11), e68, 2017; Volz & Gazzaniga, Brain, 140(7), 2051-2060, 2017; Volz, Hillyard, Miller, & Gazzaniga, Brain, 141(3), e15, 2018). This collective review paper aims to summarize the empirical common ground, to delineate the different interpretations, and to identify the remaining questions.

Visual Functions Generating Conscious Seeing.

Visual functions are reviewed that coincide with conscious as opposed to unconscious vision. Four stages of vision are identified, going from the fully invisible, to subjectively invisible, unattended, and clearly visible. It is proposed that feature extraction, categorization, and some aspects of visual inference occur during full and subjective invisibility.

Scene complexity modulates degree of feedback activity during object detection in natural scenes.

Selective brain responses to objects arise within a few hundreds of milliseconds of neural processing, suggesting that visual object recognition is mediated by rapid feed-forward activations. Yet disruption of neural responses in early visual cortex beyond feed-forward processing stages affects object recognition performance. Here, we unite these discrepant findings by reporting that object recognition involves enhanced feedback activity (recurrent processing within early visual cortex) when target objects are embedded in natural scenes that are characterized by high complexity.

Challenges for theories of consciousness: seeing or knowing, the missing ingredient and how to deal with panpsychism.

Significant progress has been made in the study of consciousness. Promising theories have been developed and a wealth of experimental data has been generated, both guiding us towards a better understanding of this complex phenomenon. However, new challenges have surfaced.

The Split-Brain Phenomenon Revisited: A Single Conscious Agent with Split Perception.

The split-brain phenomenon is caused by the surgical severing of the corpus callosum, the main route of communication between the cerebral hemispheres. The classical view of this syndrome asserts that conscious unity is abolished. The left hemisphere consciously experiences and functions independently of the right hemisphere.

Conscious visual memory with minimal attention.

Is conscious visual perception limited to the locations that a person attends? The remarkable phenomenon of change blindness, which shows that people miss nearly all unattended changes in a visual scene, suggests the answer is yes. However, change blindness is found after visual interference (a mask or a new scene), so that subjects have to rely on working memory (WM), which has limited capacity, to detect the change. Before such interference, however, a much larger capacity store, called fragile memory (FM), which is easily overwritten by newly presented visual information, is present.

Split brain: divided perception but undivided consciousness.

In extensive studies with two split-brain patients we replicate the standard finding that stimuli cannot be compared across visual half-fields, indicating that each hemisphere processes information independently of the other. Yet, crucially, we show that the canonical textbook findings that a split-brain patient can only respond to stimuli in the left visual half-field with the left hand, and to stimuli in the right visual half-field with the right hand and verbally, are not universally true. Across a wide variety of tasks, split-brain patients with a complete and radiologically confirmed transection of the corpus callosum showed full awareness of presence, and well above chance-level recognition of location, orientation and identity of stimuli throughout the entire visual field, irrespective of response type (left hand, right hand, or verbally).

The time course of natural scene perception with reduced attention.

Attention is thought to impose an informational bottleneck on vision by selecting particular information from visual scenes for enhanced processing. Behavioral evidence suggests, however, that some scene information is extracted even when attention is directed elsewhere. Here, we investigated the neural correlates of this ability by examining how attention affects electrophysiological markers of scene perception.

No-Report Paradigms: Extracting the True Neural Correlates of Consciousness.

The goal of consciousness research is to reveal the neural basis of phenomenal experience. To study phenomenology, experimenters seem obliged to ask reports from the subjects to ascertain what they experience. However, we argue that the requirement of reports has biased the search for the neural correlates of consciousness over the past decades.

Neural Correlates of Visual Short-term Memory Dissociate between Fragile and Working Memory Representations.

Evidence is accumulating that the classic two-stage model of visual STM (VSTM), comprising iconic memory (IM) and visual working memory (WM), is incomplete. A third memory stage, termed fragile VSTM (FM), seems to exist in between IM and WM [Vandenbroucke, A. R.

Expectations accelerate entry of visual stimuli into awareness.

How do expectations influence transitions between unconscious and conscious perceptual processing? According to the influential predictive processing framework, perceptual content is determined by predictive models of the causes of sensory signals. On one interpretation, conscious contents arise when predictive models are verified by matching sensory input (minimizing prediction error). On another, conscious contents arise when surprising events falsify current perceptual predictions.

Forgotten but not gone: Retro-cue costs and benefits in a double-cueing paradigm suggest multiple states in visual short-term memory.

Visual short-term memory (VSTM) performance is enhanced when the to-be-tested item is cued after encoding. This so-called retro-cue benefit is typically accompanied by a cost for the noncued items, suggesting that information is lost from VSTM upon presentation of a retrospective cue. Here we assessed whether noncued items can be restored to VSTM when made relevant again by a subsequent second cue.

Pupil size tracks perceptual content and surprise.

Changes in pupil size at constant light levels reflect the activity of neuromodulatory brainstem centers that control global brain state. These endogenously driven pupil dynamics can be synchronized with cognitive acts. For example, the pupil dilates during the spontaneous switches of perception of a constant sensory input in bistable perceptual illusions.

NMDA Receptor Antagonist Ketamine Distorts Object Recognition by Reducing Feedback to Early Visual Cortex.

It is a well-established fact that top-down processes influence neural representations in lower-level visual areas. Electrophysiological recordings in monkeys as well as theoretical models suggest that these top-down processes depend on NMDA receptor functioning. However, this underlying neural mechanism has not been tested in humans.

Top-down modulation in human visual cortex predicts the stability of a perceptual illusion.

Conscious perception sometimes fluctuates strongly, even when the sensory input is constant. For example, in motion-induced blindness (MIB), a salient visual target surrounded by a moving pattern suddenly disappears from perception, only to reappear after some variable time. Whereas such changes of perception result from fluctuations of neural activity, mounting evidence suggests that the perceptual changes, in turn, may also cause modulations of activity in several brain areas, including visual cortex.

Consciousness is not necessary for visual feature binding.

When visual information enters the brain, it is relayed to different specialized regions, processing features such as shape, color, or motion. And yet, in our conscious experience of a colored, moving shape, all the different features seem to be integrated into one unified percept. Therefore, it has been hypothesized that consciousness and feature binding share an intimate relationship.

Latent memory of unattended stimuli reactivated by practice: an FMRI study on the role of consciousness and attention in learning.

Although we can only report about what is in the focus of our attention, much more than that is actually processed. And even when attended, stimuli may not always be reportable, for instance when they are masked. A stimulus can thus be unreportable for different reasons: the absence of attention or the absence of a conscious percept.

The subjective experience of object recognition: comparing metacognition for object detection and object categorization.

Perceptual decisions seem to be made automatically and almost instantly. Constructing a unitary subjective conscious experience takes more time. For example, when trying to avoid a collision with a car on a foggy road you brake or steer away in a reflex, before realizing you were in a near accident.

Accurate metacognition for visual sensory memory representations.

The capacity to attend to multiple objects in the visual field is limited. However, introspectively, people feel that they see the whole visual world at once. Some scholars suggest that this introspective feeling is based on short-lived sensory memory representations, whereas others argue that the feeling of seeing more than can be attended to is illusory.