Neural mechanisms underlying conscious and unconscious vision. Evidence from event-related potentials and transcranial magnetic stimulation

Vision affords us with the ability to consciously see, and use this information in our behavior. While research has produced a detailed account of the function of the visual system, the neural processes that underlie conscious vision are still debated. One of the aims of the present thesis was to examine the time-course of the neuroelectrical processes that correlate with conscious vision. The second aim was to study the neural basis of unconscious vision, that is, situations where a stimulus that is not consciously perceived nevertheless influences behavior. According to current prevalent models of conscious vision, the activation of visual cortical areas is not, as such, sufficient for consciousness to emerge, although it might be sufficient for unconscious vision. Conscious vision is assumed to require reciprocal communication between cortical areas, but views differ substantially on the extent of this recurrent communication. Visual consciousness has been proposed to emerge from recurrent neural interactions within the visual system, while other models claim that more widespread cortical activation is needed for consciousness. Studies I-III compared models of conscious vision by studying event-related potentials (ERP). ERPs represent the brain’s average electrical response to stimulation. The results support the model that associates conscious vision with activity localized in the ventral visual cortex. The timing of this activity corresponds to an intermediate stage in visual processing. Earlier stages of visual processing may influence what becomes conscious, although these processes do not directly enable visual consciousness. Late processing stages, when more widespread cortical areas are activated, reflect the access to and manipulation of contents of consciousness. Studies IV and V concentrated on unconscious vision. By using transcranial magnetic stimulation (TMS) we show that when early visual cortical processing is disturbed so that subjects fail to consciously perceive visual stimuli, they may nevertheless guess (above chance-level) the location where the visual stimuli were presented. However, the results also suggest that in a similar situation, early visual cortex is necessary for both conscious and unconscious perception of chromatic information (i.e. color). Chromatic information that remains unconscious may influence behavioral responses when activity in visual cortex is not disturbed by TMS. Our results support the view that early stimulus-driven (feedforward) activation may be sufficient for unconscious processing. In conclusion, the results of this thesis support the view that conscious vision is enabled by a series of processing stages. The processes that most closely correlate with conscious vision take place in the ventral visual cortex ~200 ms after stimulus presentation, although preceding time-periods and contributions from other cortical areas such as the parietal cortex are also indispensable. Unconscious vision relies on intact early visual activation, although the location of visual stimulus may be unconsciously resolved even when activity in the early visual cortex is interfered with.Siirretty Doriast

Multiple independent components contribute to event-related potential correlates of conscious vision

Research has revealed two major event-related potential (ERP) markers of visual awareness: the earlier Visual Awareness Negativity (VAN, around 150-250 ms after stimulus onset), and the following Late Positivity (LP, around 300-500 ms after stimulus onset). Understanding the neural sources that give rise to VAN and LP is important in order to understand what kind of neural processes underlie conscious visual perception. Although the ERPs afford high temporal resolution, their spatial resolution is limited because multiple separate neural sources sum up at the scalp level. In the present study, we sought to characterize the locations and time-courses of independent neural sources underlying the ERP correlates of visual awareness by means of Independent Component Analysis (ICA). ICA allows identifying and localizing the temporal dynamics of different neural sources that contribute to the ERP correlates of conscious perception. The present results show that the cortical sources of VAN are localized to posterior areas including occipital and temporal cortex, while LP reflects a combination of multiple sources distributed among frontal, parietal and occipito-temporal cortex. Our findings suggest that conscious vision correlates with dynamically changing neural sources, developing in part in "accumulative fashion": consciousness-related activity initially arises in few early sources and, subsequently, additional sources are engaged as a function of time. The results further suggest that even early latency neural sources that correlate with conscious perception may also associate with action-related processes

Consciousness as a concrete physical phenomenon

The typical empirical approach to studying consciousness holds that we can only observe the neural correlates of experiences, not the experiences themselves. In this paper we argue, in contrast, that experiences are concrete physical phenomena that can causally interact with other phenomena, including observers. Hence, experiences can be observed and scientifically modelled. We propose that the epistemic gap between an experience and a scientific model of its neural mechanisms stems from the fact that the model is merely a theoretical construct based on observations, and distinct from the concrete phenomenon it models, namely the experience itself. In this sense, there is a gap between any natural phenomenon and its scientific model. On this approach, a neuroscientific theory of the constitutive mechanisms of an experience is literally a model of the subjective experience itself. We argue that this metatheoretical framework provides a solid basis for the empirical study of consciousness.</p

Promise and challenges for discovering transcranial magnetic stimulation induced "numbsense"-Commentary on Ro & Koenig (2021).

The notion that behavioral responses to stimuli can be mediated by separate unconscious and conscious sensory pathways remains popular, but also hotly debated. Recently, Ro and Koenig (2021) reported that when activity in somatosensory cortex was interfered with transcranial magnetic stimulation (TMS), participants could discriminate tactile stimuli they reported not consciously feeling. The study launches an interesting new area of research, helping to uncover mechanisms of unconscious perception that possibly generalize across different sensory modalities. However, we argue here that the study by Ro and Koenig also has several significant shortcomings, and it fails to provide evidence that pathways bypassing primary somatosensory cortex enable unconscious tactile discrimination. By referring to numerous studies investigating TMS-induced blindsight, we outline challenges in demonstrating unconscious sensory pathways using TMS. By facing to these challenges, research investigating TMS-induced numbsense has potential to stimulate progress in stubborn debates and reveal modality-general mechanisms of unconscious perception

Is the primary visual cortex necessary for blindsight-like behavior? Review of transcranial magnetic stimulation studies in neurologically healthy individuals

The visual pathways that bypass the primary visual cortex (V1) are often assumed to support visually guided behavior in humans in the absence of conscious vision. This conclusion is largely based on findings on patients: V1 lesions cause blindness but sometimes leave some visually guided behaviors intact-this is known as blindsight. With the aim of examining how well the findings on blindsight patients generalize to neurologically healthy individuals, we review studies which have tried to uncover transcranial magnetic stimulation (TMS) induced blindsight. In general, these studies have failed to demonstrate a completely unconscious blindsight-like capacity in neurologically healthy individuals. A possible exception to this is TMS-induced blindsight of stimulus presence or location. Because blindsight in patients is often associated with some form of introspective access to the visual stimulus, and blindsight may be associated with neural reorganization, we suggest that rather than revealing a dissociation between visually guided behavior and conscious seeing, blindsight may reflect preservation or partial recovery of conscious visual perception after the lesion

Subliminal perception is continuous with conscious vision and can be predicted from prestimulus electroencephalographic activity

Individuals are able to discriminate visual stimuli they report not consciously seeing. This phenomenon is known as "subliminal perception." Such capacity is often assumed to be relatively automatic in nature and rely on stimulus-driven activity in low-level cortical areas. Instead, here we asked to what extent neural activity before stimulus presentation influences subliminal perception. We asked participants to discriminate the location of a briefly presented low-contrast visual stimulus and then rate how well they saw the stimulus. Consistent with previous studies, participants correctly discriminated with slightly above chance-level accuracy the location of a stimulus they reported not seeing. Signal detection analyses indicated that while subjects categorized their percepts as "unconscious," their capacity to discriminate these stimuli lay on the same continuum as conscious vision. We show that the accuracy of discriminating the location of a subliminal stimulus could be predicted with relatively high accuracy (AUC = 0.70) based on lateralized electroencephalographic (EEG) activity before the stimulus, the hemifield where the stimulus was presented, and the accuracy of previous trial's discrimination response. Altogether, our results suggest that rather than being a separate unconscious capacity, subliminal perception is based on similar processes as conscious vision.</p

Binocular disparity can augment the capacity of vision without affecting subjective experience of depth

Binocular disparity results in a tangible subjective experience of three-dimensional world, but whether disparity also augments objective perceptual performance remains debated. We hypothesized that the improved coding of depth enabled by binocular disparity allows participants to individuate more objects at a glance as the objects can be more efficiently differentiated from each other and the background. We asked participants to enumerate objects in briefly presented naturalistic (Experiment 1) and artificial (Experiment 2) scenes in immersive virtual reality. This type of enumeration task yields well-documented capacity limits where up to 3-4 items can be enumerated rapidly and accurately, known as subitizing. Our results show that although binocular disparity did not yield a large general improvement in enumeration accuracy or reaction times, it improved participants' ability to process the items right after the limit of perceptual capacity. Binocular disparity also sped-up response times by 27 ms on average when artificial stimuli (cubes) were used. Interestingly, the influence of disparity on subjectively experienced depth revealed a clearly different pattern than the influence of disparity on objective performance. This suggests that the functional and subjective sides of stereopsis can be dissociated. Altogether our results suggest that binocular disparity may increase the number of items the visual system can simultaneously process. This may help animals to better resolve and track objects in complex, cluttered visual environments

Event-related potential correlates of learning to produce novel foreign phonemes

Learning to pronounce a foreign phoneme requires an individual to acquire a motor program that enables the reproduction of the new acoustic target sound. This process is largely based on the use of auditory feedback to detect pronunciation errors to adjust vocalization. While early auditory evoked neural activity underlies automatic detection and adaptation to vocalization errors, little is known about the neural correlates of acquiring novel speech targets. To investigate the neural processes that mediate the learning of foreign phoneme pronunciation, we recorded event-related potentials when participants (N 19) pronounced native or foreign phonemes. Behavioral results indicated that the participants’ pronunciation of the foreign phoneme improved during the experiment. Early auditory responses (N1 and P2 waves, approximately 85–290 ms after the sound onset) revealed no differences between foreign and native phonemes. In contrast, the amplitude of the frontocentrally distributed late slow wave (LSW, 320–440 ms) was modulated by the pronunciation of the foreign phonemes, and the effect changed during the experiment, paralleling the improvement in pronunciation. These results suggest that the LSW may reflect higher-order monitoring processes that signal successful pronunciation and help learn novel phonemes.</p

Modality-specific and modality-general electrophysiological correlates of visual and auditory awareness: Evidence from a bimodal ERP experiment

To date, most studies on the event-related potential (ERP) correlates of conscious perception have examined a single perceptual modality. We compared electrophysiological correlates of visual and auditory awareness in the same experiment to test whether there are modality-specific and modality-general correlates of conscious perception. We used near threshold stimulation and analyzed event-related potentials in response to aware and unaware trials in visual, auditory and bimodal conditions. The results showed modality-specific negative amplitude correlates of conscious perception between 200 and 300 ms after stimulus onset. A combination of these auditory and visual awareness negativities was observed in the bimodal condition. A later positive amplitude difference, whose early part was modality-specific, possibly reflecting access to global workspace, and later part shared modality-general features, possibly indicating higher level cognitive processing involving the decision making, was also observed

Segregated brain state during hypnosis

Can the brain be shifted into a different state using a simple social cue, as tests on highly hypnotizable subjects would suggest? Demonstrating an altered global brain state is difficult. Brain activation varies greatly during wakefulness and can be voluntarily influenced. We measured the complexity of electrophysiological response to transcranial magnetic stimulation in one ‘hypnotic virtuoso’. Such a measure produces a response arguably outside the subject’s voluntary control and has been proven adequate for discriminating conscious from unconscious brain states. We show that a single-word hypnotic induction robustly shifted global neural connectivity into a state where activity remained sustained but failed to ignite strong, coherent activity in frontoparietal cortices. Changes in perturbational complexity indicate a similar move towards a more segregated state. We interpret these findings to suggest a shift in the underlying state of the brain, likely moderating subsequent hypnotic responding.</p