Visual Consciousness Research Articles

Dissociations of conscious and unconscious perception in TMS-induced blindsight

As with some patients with primary visual cortex (V1) damage, transcranial magnetic stimulation (TMS) over V1 reliably induces blindsight, whereby observers can correctly discriminate the attributes of visual stimuli despite being unable to detect them. This TMS-induced blindsight has been demonstrated to reflect a form of unconscious vision that relies upon different neural pathways than with conscious vision. However, the timing of the neural processes mediating TMS-induced blindsight has been unclear, especially when considering suggestions that TMS interferes with feedback processes to V1 that mediate conscious visual perception. To better elucidate the neural mechanisms that give rise to blindsight, we tested TMS-induced blindsight for the orientation of visual stimuli across a range of stimulus onset asynchronies (SOAs) to assess how different latencies of visual cortex disruption, relative to a visual stimulus, affect detection rates and forced-choice discrimination accuracy. At all TMS latencies, including at SOAs with substantial visual suppression from TMS, discrimination performance was significantly above-chance, demonstrating the consistency of TMS-induced blindsight. Crucially, we observed two windows of maximum visual suppression from TMS at SOAs between 65 and 105 ms, but consistent above-chance discrimination performance accuracy across these windows. However, at longer SOAs, detection and discrimination covaried, suggesting a dependency of discrimination performance on detection only when detection rates exceed threshold levels of normal vision. Taken together, these results indicate that unconscious discrimination occurs independently of detection, including at TMS intervals that optimally interfere with conscious visual perception. They further suggest that forced-choice discrimination is less dependent on feedback processes to V1 than visual awareness and that TMS-induced blindsight is not the same as near-threshold vision.

Robust Visual Responses and Normal Retinotopy in Primate Lateral Geniculate Nucleus following Long-term Lesions of Striate Cortex.

Lesions of striate cortex (V1) trigger massive retrograde degeneration of neurons in the LGN. In primates, these lesions also lead to scotomas, within which conscious vision is abolished. Mediation of residual visual capacity within these regions (blindsight) has been traditionally attributed to an indirect visual pathway to the extrastriate cortex, which involves the superior colliculus and pulvinar complex. However, recent studies have suggested that preservation of the LGN is critical for behavioral evidence of blindsight, raising the question of what type of visual information is channeled by remaining neurons in this structure. A possible contribution of LGN neurons to blindsight is predicated on two conditions: that the neurons that survive degeneration remain visually responsive, and that their receptive fields continue to represent the region of the visual field inside the scotoma. We tested these conditions in male and female marmoset monkeys (Callithrix jacchus) with partial V1 lesions at three developmental stages (early postnatal life, young adulthood, old age), followed by long recovery periods. In all cases, recordings from the degenerated LGN revealed neurons with well-formed receptive fields throughout the scotoma. The responses were consistent and robust, and followed the expected eye dominance and retinotopy observed in the normal LGN. The responses had short latencies and preceded those of neurons recorded in the extrastriate middle temporal area. These findings suggest that the pathway that links LGN neurons to the extrastriate cortex is physiologically viable and can support residual vision in animals with V1 lesions incurred at various ages.SIGNIFICANCE STATEMENT Patients with a lesion of the primary visual cortex (V1) can retain certain visually mediated behaviors, particularly if the lesion occurs early in life. This phenomenon ("blindsight") not only sheds light on the nature of consciousness, but also has implications for studies of brain circuitry, development, and plasticity. However, the pathways that mediate blindsight have been the subject of debate. Recent studies suggest that projections from the LGN might be critical, but this finding is puzzling given that the lesions causes severe cell death in the LGN. Here we demonstrate in monkeys that the surviving LGN neurons retain a remarkable level of visual function and could therefore be the source of the visual information that supports blindsight.

“The Progress of Painting”: Leigh Hunt’s Progress of Visual Sense

This essay examines the progress of Leigh Hunt’s visual sense to reconsider his cultural identity within the wider context of the metropolitan visual culture of the Romantic era. By paying close attention to Hunt’s visual experiences, especially during his adolescence, this essay argues that the juvenile Hunt’s strong admiration for the visual certainly affected the novel attempt to realize his ekphrastic aspiration with a pictorial method, as has been actively discussed by recent critics. First, along with a close re-reading of primary sources, including Hunt’s autobiography and letters, the essay examines how the schoolboy Hunt developed his visual consciousness inside and outside of Christ’s Hospital School. Subsequently, reading Hunt’s adolescent work, “The Progress of Painting” from Juvenilia, the essay especially focuses on the aesthetic interrelationship between Hunt and West, who affected the young Hunt’s visual knowledge of art and ekphrastic interpretation. Although there are limitations in attempting to fully decipher the young Hunt’s visual sense, this essay can be a cornerstone to reconsider the aesthetic value of Hunt and his literary works in relation to Romantic visual culture.

Insula mediates heartbeat related effects on visual consciousness

Interoceptive signals, such as the heartbeat, are processed in a network of brain regions including the insular cortex. Recent studies have shown that such signals modulate perceptual and cognitive processing, and that they impact visual awareness. For example, visual stimuli presented synchronously to the heartbeat take longer to enter visual awareness than the same stimuli presented asynchronously to the heartbeat, and this is reflected in anterior insular activation. This finding demonstrated a link between the processing of interoceptive and exteroceptive signals as well as visual awareness in the insular cortex. The advantage for visual stimuli which are asynchronous to the heartbeat to enter visual consciousness may indicate a role for the anterior insula in the suppression of the sensory consequences of cardiac signals. Here, we present data from the detailed investigation of two patients with insular lesions (as well as four patients with non-insular lesions and healthy age matched controls) indicating that a lesion of the anterior insular cortex, but not of other regions, abolished this cardio-visual suppression effect. The present data provide causal evidence for the role of the anterior insula in the integration of internal interoceptive and external sensory signals for visual awareness.

Ensemble Perception.

To understand visual consciousness, we must understand how the brain represents ensembles of objects at many levels of perceptual analysis. Ensemble perception refers to the visual system's ability to extract summary statistical information from groups of similar objects-often in a brief glance. It defines foundational limits on cognition, memory, and behavior. In this review, we provide an operational definition of ensemble perception and demonstrate that ensemble perception spans across multiple levels of visual analysis, incorporating both low-level visual features and high-level social information. Further, we investigate the functional usefulness of ensemble perception and its efficiency, and we consider possible physiological and cognitive mechanisms that underlie an individual's ability to make accurate and rapid assessments of crowds of objects.

The structure of sensorimotor explanation

The sensorimotor theory of vision and visual consciousness is often described as a radical alternative to the computational and connectionist orthodoxy in the study of visual perception. However, it is far from clear whether the theory represents a significant departure from orthodox approaches or whether it is an enrichment of it. In this study, I tackle this issue by focusing on the explanatory structure of the sensorimotor theory. I argue that the standard formulation of the theory subscribes to the same theses of the dynamical hypothesis and that it affords covering-law explanations. This however exposes the theory to the mere description worry and generates a puzzle about the role of representations. I then argue that the sensorimotor theory is compatible with a mechanistic framework, and show how this can overcome the mere description worry and solve the problem of the explanatory role of representations. By doing so, it will be shown that the theory should be understood as an enrichment of the orthodoxy, rather than an alternative.

Do early neural correlates of visual consciousness show the oblique effect? A binocular rivalry and event-related potential study.

When dissimilar images are presented one to each eye, we do not see both images; rather, we see one at a time, alternating unpredictably. This is called binocular rivalry, and it has recently been used to study brain processes that correlate with visual consciousness, because perception changes without any change in the sensory input. Such studies have used various types of images, but the most popular have been gratings: sets of bright and dark lines of orthogonal orientations presented one to each eye. We studied whether using cardinal rival gratings (vertical, 0°, and horizontal, 90°) versus oblique rival gratings (left-oblique, –45°, and right-oblique, 45°) influences early neural correlates of visual consciousness, because of the oblique effect: the tendency for visual performance to be greater for cardinal gratings than for oblique gratings. Participants viewed rival gratings and pressed keys indicating which of the two gratings they perceived, was dominant. Next, we changed one of the gratings to match the grating shown to the other eye, yielding binocular fusion. Participants perceived the rivalry-to-fusion change to the dominant grating and not to the other, suppressed grating. Using event-related potentials (ERPs), we found neural correlates of visual consciousness at the P1 for both sets of gratings, as well as at the P1-N1 for oblique gratings, and we found a neural correlate of the oblique effect at the N1, but only for perceived changes. These results show that the P1 is the earliest neural activity associated with visual consciousness and that visual consciousness might be necessary to elicit the oblique effect.

Montage October: Dialectic of the Shot

In “Montage October,” which appeared in Artforum in January 1973, Rosalind Krauss argues that Sergei Eisenstein's 1928 film was not only a celebration of a Marxist victory but also a demonstration of Marxist dialectical reasoning through montage. Krauss contends that rather than passively reconstructing the chain of circumstances that culminated in the storming of the Winter Palace in 1917, Eisenstein hoped to draw a filmic equivalence between the leap of revolutionary consciousness, which opens up access to the future by transcending the real, and the leap of visual consciousness, which goes beyond the normal bounds of filmic space.

Binocular Rivalry: A Window into Cortical Competition and Suppression

When the two eyes view very dissimilar images, the visual system often fails to combine the images and one experiences stochastically alternating percepts. This phenomenon, called binocular rivalry, has fascinated researchers for centuries since it provides insights into two critical aspects of visual perception: visual consciousness and cortical suppression. Here, we review the mechanisms of binocular rivalry from a cognitive neuroscience perspective, focusing on empirical findings from two widely used methods—functional magnetic resonance imaging (fMRI) and electroencephalography (EEG). With these techniques, researchers have been able to identify the cortical sites of suppression in binocular rivalry, probe neural responses evoked by unconscious (invisible) visual stimuli, and examine the role of top-down attentional signals in rivalry. We conclude by proposing some future directions for the study of binocular rivalry.

More than blindsight: Case report of a child with extraordinary visual capacity following perinatal bilateral occipital lobe injury

Injury to the primary visual cortex (V1, striate cortex) and the geniculostriate pathway in adults results in cortical blindness, abolishing conscious visual perception. Early studies by Larry Weiskrantz and colleagues demonstrated that some patients with an occipital-lobe injury exhibited a degree of unconscious vision and visually-guided behaviour within the blind field. A more recent focus has been the observed phenomenon whereby early-life injury to V1 often results in the preservation of visual perception in both monkeys and humans. These findings initiated a concerted effort on multiple fronts, including nonhuman primate studies, to uncover the neural substrate/s of the spared conscious vision. In both adult and early-life cases of V1 injury, evidence suggests the involvement of the Middle Temporal area (MT) of the extrastriate visual cortex, which is an integral component area of the dorsal stream and is also associated with visually-guided behaviors. Because of the limited number of early-life V1 injury cases for humans, the outstanding question in the field is what secondary visual pathways are responsible for this extraordinary capacity? Here we report for the first time a case of a child (B.I.) who suffered a bilateral occipital-lobe injury in the first two weeks postnatally due to medium-chain acyl-Co-A dehydrogenase deficiency. At 6 years of age, B.I. underwent a battery of neurophysiological tests, as well as structural and diffusion MRI and ophthalmic examination at 7 years. Despite the extensive bilateral occipital cortical damage, B.I. has extensive conscious visual abilities, is not blind, and can use vision to navigate his environment. Furthermore, unlike blindsight patients, he can readily and consciously identify happy and neutral faces and colors, tasks associated with ventral stream processing. These findings suggest significant re-routing of visual information. To identify the putative visual pathway/s responsible for this ability, MRI tractography of secondary visual pathways connecting MT with the lateral geniculate nucleus (LGN) and the inferior pulvinar (PI) were analysed. Results revealed an increased PI-MT pathway in the left hemisphere, suggesting that this pulvinar relay could be the neural pathway affording the preserved visual capacity following an early-life lesion of V1. These findings corroborate anatomical evidence from monkeys showing an enhanced PI-MT pathway following an early-life lesion of V1, compared to adults.

Dissociable Electroencephalograph Correlates of Visual Awareness and Feature-Based Attention.

Background: The relationship between awareness and attention is complex and controversial. A growing body of literature has shown that the neural bases of consciousness and endogenous attention (voluntary attention) are independent. The important role of exogenous attention (reflexive attention) on conscious experience has been noted in several studies. However, exogenous attention can also modulate subliminal processing, suggesting independence between the two processes. The question of whether visual awareness and exogenous attention rely on independent mechanisms under certain circumstances remains unanswered.Methods: In the current study, electroencephalograph recordings were conducted using 64 channels from 16 subjects while subjects attempted to detect faint speed changes of colored rotating dots. Awareness and attention were manipulated throughout trials in order to test whether exogenous attention and visual awareness rely on independent mechanisms.Results: Neural activity related to consciousness was recorded in the following cue-locked time-windows (event related potential, cluster- based permutation test): 0–50, 150–200, and 750–800 ms. With a more liberal threshold, the inferior occipital lobe was found to be the source of awareness-related activity in the 0–50 ms range. In the later 150–200 ms range, activity in the fusiform and post-central gyrus was related to awareness. Awareness-related activation in the later 750–800 ms range was more widely distributed. This awareness-related activation pattern was quite different from that of attention. Attention-related neural activity was emphasized in the 750–800 ms time window and the main source of attention-related activity was localized to the right angular gyrus. These results suggest that exogenous attention and visual consciousness correspond to different and relatively independent neural mechanisms and are distinct processes under certain conditions.

Micro-calibration of space and motion by photoreceptors synchronized in parallel with cortical oscillations: A unified theory of visual perception

A fundamental function of the visual system is detecting motion, yet visual perception is poorly understood. Current research has determined that the retina and ganglion cells elicit responses for motion detection; however, the underlying mechanism for this is incompletely understood. Previously we proposed that retinogeniculo-cortical oscillations and photoreceptors work in parallel to process vision. Here we propose that motion could also be processed within the retina, and not in the brain as current theory suggests. In this paper, we discuss: 1) internal neural space formation; 2) primary, secondary, and tertiary roles of vision; 3) gamma as the secondary role; and 4) synchronization and coherence. Movement within the external field is instantly detected by primary processing within the space formed by the retina, providing a unified view of the world from an internal point of view. Our new theory begins to answer questions about: 1) perception of space, erect images, and motion, 2) purpose of lateral inhibition, 3) speed of visual perception, and 4) how peripheral color vision occurs without a large population of cones located peripherally in the retina. We explain that strong oscillatory activity influences on brain activity and is necessary for: 1) visual processing, and 2) formation of the internal visuospatial area necessary for visual consciousness, which could allow rods to receive precise visual and visuospatial information, while retinal waves could link the lateral geniculate body with the cortex to form a neural space formed by membrane potential-based oscillations and photoreceptors. We propose that vision is tripartite, with three components that allow a person to make sense of the world, terming them “primary, secondary, and tertiary roles” of vision. Finally, we propose that Gamma waves that are higher in strength and volume allow communication among the retina, thalamus, and various areas of the cortex, and synchronization brings cortical faculties to the retina, while the thalamus is the link that couples the retina to the rest of the brain through activity by gamma oscillations. This novel theory lays groundwork for further research by providing a theoretical understanding that expands upon the functions of the retina, photoreceptors, and retinal plexus to include parallel processing needed to form the internal visual space that we perceive as the external world.

Does TMS on V3 block conscious visual perception?

Primary visual cortex (V1) and extrastriate V2 are necessary for the emergence of visual consciousness, but the effects of involvement of extrastriate V3 on visual consciousness is unclear. The objective of this study was to examine the causal role of V3 in visual consciousness in humans. We combined neuronavigated transcranial magnetic stimulation (TMS) with a computational model of the TMS-induced electric field to test whether or not the intact processing of visual input in V3, like in V1 and V2, is necessary for conscious visual perception. We targeted the stimulation both to V2 and to V3. If TMS of V3 blocks conscious visual perception of stimuli, then activation in V3 is a causally necessary prerequisite for conscious perception of stimuli. According to the alternative hypothesis, TMS of V3 will not block the conscious visual perception of stimuli, because the pathways from V1 to the higher cortical areas that go around V3 provide sufficient visual input for the emergence of conscious visual perception. The results showed that TMS interfered with conscious perception of features, detection of stimulus presence and the ability to discriminate the letter stimuli both when TMS was targeted either to V3 or to V2. For the conscious detection of stimulus presence, the effect was significantly stronger when V2 was stimulated than when V3 was stimulated. The results of the present study suggest that in addition to the primary visual cortex and V2, also V3 causally contributes to the generation of the most basic form of visual consciousness. Importantly, the results also indicate that V3 is necessary for visual perception in general, not only for visual consciousness.

On the “blindness” of blindsight: What is the evidence for phenomenal awareness in the absence of primary visual cortex (V1)?

Blindsight has been central to theories of phenomenal awareness; that a lesion to primary visual cortex (V1) abolishes all phenomenal awareness while unconscious visual functions can remain has led to the view that this region plays a crucial role in generating visual consciousness. However, since the early 20th century, there have been reports, many of which controversial, of phenomenal awareness in patients with V1 lesions. These reports include selective sparing of motion awareness, hemianopic completion and visual aftereffects. More recently, there have been successful attempts of inducing visual qualia with noninvasive brain stimulation. Here we critically review this evidence and discuss their implications to theoretical understanding of phenomenal awareness.

Visually evoked responses from the blind field of hemianopic patients

Hemianopia is a visual field defect characterized by decreased vision or blindness in the contralesional visual field of both eyes. The presence of well documented above-chance unconscious behavioural responses to visual stimuli presented to the blind hemifield (blindsight) has stimulated a great deal of research on the neural basis of this important phenomenon. The present study is concerned with electrophysiological responses from the blind field. Since previous studies found that transient Visual Evoked Potentials (VEPs) are not entirely suitable for this purpose here we propose to use Steady-State VEPs (SSVEPs). A positive result would have important implications for the understanding of the neural bases of conscious vision. We carried out a passive SSVEP stimulation with healthy participants and hemianopic patients. Stimuli consisted of four black-and-white sinusoidal Gabor gratings presented one in each visual field quadrant and flickering one at a time at a 12Hz rate. To assess response reliability a Signal-to-Noise Ratio analysis was conducted together with further analyses in time and frequency domains to make comparisons between groups (healthy participants and patients), side of brain lesion (left and right) and visual fields (sighted and blind).The important overall result was that stimulus presentation to the blind hemifield yielded highly reliable responses with time and frequency features broadly similar to those found for cortical extrastriate areas in healthy controls. Moreover, in the intact hemifield of hemianopics and in healthy controls there was evidence of a role of prefrontal structures in perceptual awareness. Finally, the presence of different patterns of brain reorganization depended upon the side of lesion.

Can perceptual grouping unfold in the absence of visual consciousness?

Can perceptual grouping unfold in the absence of visual consciousness?

From Cortical Blindness to Conscious Visual Perception: Theories on Neuronal Networks and Visual Training Strategies.

Homonymous hemianopia (HH) is the most common cortical visual impairment leading to blindness in the contralateral hemifield. It is associated with many inconveniences and daily restrictions such as exploration and visual orientation difficulties. However, patients with HH can preserve the remarkable ability to unconsciously perceive visual stimuli presented in their blindfield, a phenomenon known as blindsight. Unfortunately, the nature of this captivating residual ability is still misunderstood and the rehabilitation strategies in terms of visual training have been insufficiently exploited. This article discusses type I and type II blindsight in a neuronal framework of altered global workspace, resulting from inefficient perception, attention and conscious networks. To enhance synchronization and create global availability for residual abilities to reach visual consciousness, rehabilitation tools need to stimulate subcortical extrastriate pathways through V5/MT. Multisensory bottom-up compensation combined with top-down restitution training could target pre-existing and new neuronal mechanisms to recreate a framework for potential functionality.

The Perception‐Action Model: Counting Computational Mechanisms

AbstractMilner and Goodale's Two Visual Systems Hypothesis (TVSH) is regarded as common ground in recent discussions of visual consciousness. A central part of TVSH is a functional model of vision and action (a functional perception‐action model, PAM for short). In this paper, I provide a brief overview of these current discussions and argue that there is ambiguity between a strong and a weak version of PAM. I argue that, given a standard way of individuating computational mechanisms, the available evidence cannot be used to distinguish between these versions. This not only has consequences for philosophical theories of the role of visual consciousness but also for the role of experimental evidence in model testing in cognitive neuroscience.

Oscillatory Correlates of Visual Consciousness

Conscious experiences are linked to activity in our brain: the neural correlates of consciousness (NCC). Empirical research on these NCCs covers a wide range of brain activity signals, measures, and methodologies. In this paper, we focus on spontaneous brain oscillations; rhythmic fluctuations of neuronal (population) activity which can be characterized by a range of parameters, such as frequency, amplitude (power), and phase. We provide an overview of oscillatory measures that appear to correlate with conscious perception. We also discuss how increasingly sophisticated techniques allow us to study the causal role of oscillatory activity in conscious perception (i.e., ‘entrainment’). This review of oscillatory correlates of consciousness suggests that, for example, activity in the alpha-band (7–13 Hz) may index, or even causally support, conscious perception. But such results also showcase an increasingly acknowledged difficulty in NCC research; the challenge of separating neural activity necessary for conscious experience to arise (prerequisites) from neural activity underlying the conscious experience itself (substrates) or its results (consequences).

A subset of ipRGCs regulates both maturation of the circadian clock and segregation of retinogeniculate projections in mice.

The visual system consists of two major subsystems, image-forming circuits that drive conscious vision and non-image-forming circuits for behaviors such as circadian photoentrainment. While historically considered non-overlapping, recent evidence has uncovered crosstalk between these subsystems. Here, we investigated shared developmental mechanisms. We revealed an unprecedented role for light in the maturation of the circadian clock and discovered that intrinsically photosensitive retinal ganglion cells (ipRGCs) are critical for this refinement process. In addition, ipRGCs regulate retinal waves independent of light, and developmental ablation of a subset of ipRGCs disrupts eye-specific segregation of retinogeniculate projections. Specifically, a subset of ipRGCs, comprising ~200 cells and which project intraretinally and to circadian centers in the brain, are sufficient to mediate both of these developmental processes. Thus, this subset of ipRGCs constitute a shared node in the neural networks that mediate light-dependent maturation of the circadian clock and light-independent refinement of retinogeniculate projections.