Why is “blindsight” blind? A new perspective on primary visual cortex,recurrent activity and visual awareness

The neuropsychological phenomenon of blindsight has been taken to suggest that the primary visual cortex (V1) plays a unique role in visual awareness, and that extrastriate activation needs to be fed back to V1 in order for the content of that activation to be consciously perceived. The aim of this review is to evaluate this theoretical framework and to revisit its key tenets. Firstly, is blindsight truly a dissociation of awareness and visual detection? Secondly, is there sufficient evidence to rule out the possibility that the loss of awareness resulting from a V1 lesion simply reflects reduced extrastriate responsiveness, rather than a unique role of V1 in conscious experience? Evaluation of these arguments and the empirical evidence leads to the conclusion that the loss of phenomenal awareness in blindsight may not be due to feedback activity in V1 being the hallmark awareness. On the basis of existing literature, an alternative explanation of blindsight is proposed. In this view, visual awareness is a “global” cognitive function as its hallmark is the availability of information to a large number of perceptual and cognitive systems; this requires inter-areal long-range synchronous oscillatory activity. For these oscillations to arise, a specific temporal profile of neuronal activity is required, which is established through recurrent feedback activity involving V1 and the extrastriate cortex. When V1 is lesioned, the loss of recurrent activity prevents inter-areal networks on the basis of oscillatory activity. However, as limited amount of input can reach extrastriate cortex and some extrastriate neuronal selectivity is preserved, computations involving comparison of neural firing rates within a cortical area remain possible. This enables “local” read-out from specific brain regions, allowing for the detection and discrimination of basic visual attributes. Thus blindsight is blind due to lack of “global” long-range synchrony, and it functions via “local” neural readout from extrastriate areas.     

Are there unconscious perceptual processes?

Blindsight and vision for action seem to be exemplars of unconscious visual processes. However, researchers have recently argued that blindsight is not really a kind of unconscious vision but is rather severely degraded conscious vision. Morten Overgaard and colleagues have recently developed new methods for measuring the visibility of visual stimuli. Studies using these methods show that reported clarity of visual stimuli correlates with accuracy in both normal individuals and blindsight patients. Vision for action has also come under scrutiny. Recent findings seem to show that information processed by the dorsal stream for online action contributes to visual awareness. Some interpret these results as showing that some dorsal stream processes are conscious visual processes (e.g., Gallese, 2007; Jacob & Jeannerod, 2003). The aim of this paper is to provide new support for the more traditional view that blindsight and vision for action are genuinely unconscious perceptual processes. I argue that individuals with blindsight do not have access to the kind of purely qualitative color and size information which normal individuals do. So, even though people with blindsight have a kind of cognitive consciousness, visual information processing in blindsight patients is not associated with a distinctly visual phenomenology. I argue further that while dorsal stream processing seems to contribute to visual awareness, only information processed by the early dorsal stream (V1, V2, and V3) is broadcast to working memory. Information processed by later parts of the dorsal stream (the parietal lobe) never reaches working memory and hence does not correlate with phenomenal awareness. I conclude that both blindsight and vision for action are genuinely unconscious visual processes.     

The case for characterising type-2 blindsight as a genuinely visual phenomenon

Type-2 blindsight is often characterised as involving a non-visual form of awareness that blindsight subjects experience under certain presentation conditions. This paper evaluates the claim that type-2 awareness is non-visual and the proposal that it is a cognitive form of awareness. It is argued that, contrary to the standard account, type-2 awareness is best characterised as visual both because it satisfies certain criteria for being visual and because it can accommodate facts about the phenomenon that the cognitive account cannot. The conclusion is made that type-2 blindsight is best characterised as involving a form of abnormal, degraded visual awareness.     

Blindsight: the role of feedforward and feedback corticocortical connections

When human subjects suffer from a lesion to the primary visual cortex, they lose all visual percepts in the region of space that corresponds to the site of the lesion. However, they are still capable of responding to stimuli in this region when asked to 'guess' or to execute forced-choice motor commands related to these stimuli. This phenomenon, termed blindsight, is still only partly understood. Here, the possible roles of feedforward and feedback corticocortical connections in the visual brain in the understanding of blindsight are reviewed. What emerges is substantial evidence in favor of the theory that unconscious visuo-motor transformations, as in blindsight, may be executed in an entirely feedforward processing cycle, while visual awareness is critically dependent on feedback connections to the primary visual cortex.     

Masked blindsight in normal observers: Measuring subjective and objective responses to two features of each stimulus

Recent visual masking studies that have measured visual awareness with graded subjective scales have often failed the show any evidence for unconscious visual processing in normal observers in a paradigm similar to that used in studies on blindsight patients. Without any reported awareness of the target, normal observers typically cannot discriminate target’s features better than chance. The present study examined processing of color and orientation by measuring graded awareness and forced-choice discriminations for both features in each trial. When no awareness for either feature was reported, discrimination of each feature succeed better than expected by chance, even when the other feature was incorrectly discriminated in the same trial. However, the characteristics of the mask determined whether or not masked blindsight was observed. We conclude that when the processing channels are free from intra-channel interference, unbound or weakly bound features can guide behaviour without any reported awareness in normal observers.     

Non-visual consciousness and visual images in blindsight

In a recent response paper to Brogaard (2011a), Morten Overgaard and Thor Grünbaum argue that my case for the claim that blindsight subjects are not visually conscious of the stimuli they correctly identify rests on a mistaken necessary criterion for determining whether a conscious experience is visual or non-visual. Here I elaborate on the earlier argument while conceding that the question of whether blindsight subjects are visually conscious of the visual stimuli they correctly identify largely is an empirical question. I conclude by sketching a method for testing whether blindsight subjects have visual consciousness of stimuli presented to them in their blind field.     

Absence of advantageous wagering does not mean that awareness is fully abolished

Post-decision wagering (Persaud, McLeod, & Cowey, 2007) has been proposed as a method of demonstrating that perception can occur without conscious awareness. When wagering is independent from above-chance performance there is evidence of a lack of awareness of the correctness of the first-order discriminations. However, there are reasons to believe that the contingency analysis conducted by Persaud and colleagues failed to measure “the zero accuracy-wagering criterion”. The author shows that a Pearson chi-square test employed by Persaud and colleagues is unable to accommodate the hypothesis of partial independence between accuracy and advantageous wagering. On the contrary, the problem of “the zero accuracy-wagering criterion” is best conducted with log-linear models. Moreover, log-linear analysis suggests that absence of advantageous wagering is not sufficient evidence for a lack of conscious awareness. In the case of the blindsight study, there was compelling evidence that the patient GY was partially aware of having information about the correctness of his decisions.     

Reconciling current approaches to blindsight

After decades of research, blindsight is still a mysterious and controversial topic in consciousness research. Currently, many researchers tend to think of it as an ideal phenomenon to investigate neural correlates of consciousness, whereas others believe that blindsight is in fact a kind of degraded vision rather than “truly blind”. This article considers both perspectives and finds that both have difficulties understanding all existing evidence about blindsight. In order to reconcile the perspectives, we suggest two specific criteria for a good model of blindsight, able to encompass all evidence. We propose that the REF-CON model (Overgaard & Mogensen, 2014) may work as such a model.     

Perceptual size discrimination requires awareness and late visual areas: A continuous flash suppression and interocular transfer study

We applied continuous flash suppression (CFS) during an interocular transfer paradigm to evaluate the importance of awareness and the contribution of early versus late visual structures in size recognition. Specifically, we tested if size judgements of a visible target could be influenced by a congruent or incongruent prime presented to the same or different eye. Without CFS, participants categorised a target as “small” or “large” more quickly when it was preceded by a congruent prime – regardless of whether the prime and target were presented to the same or different eye. Interocular transfer enabled us to infer that the observed priming was mediated by late visual areas. In contrast, there was no priming under CFS, which underscores the importance of awareness. We conclude that awareness and late visual structures are important for size perception and that any subconscious processing of the stimulus has minimal effect on size recognition.     

Appraising Loftus and Palmer (1974) post-event information versus concurrent commentary in the context of sport

Two experiments were conducted to examine framing effects in sport. In Experiment 1, a conceptual replication [Loftus, E. F., & Palmer, J. C. (1974). Reconstruction of automobile destruction: An example of the interaction between language and memory. Journal of Verbal Learning and Verbal Behavior, 13(5), 585–589], participants watched a hockey collision, with the hit described later in a written format as a “contact”, “bump”, or “smash”. This manipulation resulted in no differences in participants’ report of how fast the players were skating, their intentions, and the outcome of the hit. In Experiment 2, participants watched the same video clip with ongoing commentary. Those who heard the announcer describing the event as “contact” estimated a higher skating speed than participants who were exposed to the “smash” commentary. Participants who were exposed to the “bump” commentary rated the repercussions of the collision as less severe than did those exposed to the other commentaries. These findings show that the perception of magnitude hierarchy may be domain specific and suggest future avenues for exploring framing effects when one is exposed to visual stimuli.     

Experience-guided attention: Uniform and implicit

Frequently finding a visual search target in one region of space induces a spatial attentional bias toward that region. Past studies on this effect typically tested fewer than 20 participants. The small sample prevents an investigation of two properties of learning: visual field uniformity and role of explicit awareness. Pooling data from multiple studies, here we examined location probability learning from ~120,000 visual search trials across 420 participants. Participants performed a serial search task. Unbeknownst to them, the target was disproportionately likely to appear in one visual quadrant. Location probability learning (LPL) was measured as the difference in reaction time to targets in the high-probability “rich” quadrant and the low-probability “sparse” quadrants. Results showed a lack of visual field effect. LPL was equivalent for “rich” quadrant in the upper left, upper right, lower left, and lower right. Learning did not induce a hotspot diagonal to the “rich” quadrant. To the contrary, RT was the longest in the diagonal quadrant. Recognition rate of the “rich” quadrant was above chance. However, recognition accuracy was unrelated to the size of LPL. Implicit learning induces visual-field-independent changes in spatial attention.     

Transient increase of intact visual field size by high-frequency narrow-band stimulation

Three patients with visual field defects were stimulated with a square matrix pattern, either static, or flickering at frequencies that had been found to either promote or not promote blindsight performance. Comparison between pre- and post-stimulation perimetric maps revealed an increase in the size of the intact visual field but only for flicker frequencies previously found to promote blindsight. These changes were temporary but dramatic – in two instances the intact field was increased by an area of ∼30 deg² of visual angle. These results indicate that not only does specific high-frequency stimulus flicker promote blindsight, but that intact visual field size may be increased by stimulation at the same frequencies. Our findings inform speculation on both the brain mechanisms and the potency of temporal modulation for altering the functional visual field.     

Relative blindsight arises from a criterion confound in metacontrast masking: implications for theories of consciousness

Relative blindsight is said to occur when different levels of subjective awareness are obtained at equality of objective performance. Using metacontrast masking, Lau and Passingham (2006) reported relative blindsight in normal observers at the shorter of two stimulus-onset asynchronies (SOAs) between target and mask. Experiment 1 replicated the critical asymmetry in subjective awareness at equality of objective performance. We argue that this asymmetry cannot be regarded as evidence for relative blindsight because the observers' responses were based on different attributes of the stimuli (criterion contents) at the two SOAs. With an invariant criterion content (Experiment 2), there was no asymmetry in subjective awareness across the two SOAs even though objective performance was the same. Experiment 3 examined the effect of criterion level on estimates of relative blindsight. Collectively, the present results question whether metacontrast masking is a suitable paradigm for establishing relative blindsight. Implications for theories of consciousness are discussed.     

Visual awareness and the levels of processing hypothesis: A critical review

Does a visual percept emerge to consciousness in a graded manner (i.e. evolving through increasing degrees of clarity), or according to a dichotomous, “all-or-none” pattern (i.e. abruptly transitioning from unawareness to awareness)? The level of processing hypothesis (LoP; B. Windey and A. Cleeremans, 2015) recently proposed a theoretical framework where the transition from unaware to aware visual experience is graded for low-level stimulus representations (i.e. stimulus “energy” or “feature” levels) whereas it is dichotomous for high-level (i.e. the perception of “letters”, “words” or “meaning”) stimulus perception. Here, we will critically review current behavioral and brain-based evidence on the LoP hypothesis and discuss potential challenges (such as differences in LoP conceptualizations, awareness scale related issues, attentional confounds and divergences on experimental factors or statistical analyses) which might be of use for future research within the field. Overall, the LoP hypothesis is a recent and promising proposal that attempts to integrate divergent evidence on the graded vs. dichotomous emergence of awareness debate. Whereas current evidence validates some of the assumptions proposed by the LoP account, there is still much work to do on both methodological and experimental levels. Future neuroimaging studies might help to disentangle the current complex pattern of results found in LoP studies and, importantly, shed some light on the ongoing debate about the search for the Neural Correlates of Consciousness (NCC).     

What is it like to have type-2 blindsight? Drawing inferences from residual function in type-1 blindsight

Controversy surrounds the question of whether the experience sometimes elicited by visual stimuli in blindsight (type-2 blindsight) is visual in nature or whether it is some sort of non-visual experience. The suggestion that the experience is visual seems, at face value, to make sense. I argue here, however, that the residual abilities found in type-1 blindsight (blindsight in which stimuli elicit no conscious experience) are not aspects of normal vision with consciousness deleted, but are based fragments of visual processes that, in themselves, would not be intelligible as visual experiences. If type-2 blindsight is a conscious manifestation of this residual function then it is not obvious that type-2 blindsight would be phenomenally like vision.     

The Transference is Indeed a Cross: Commentary on Paper by Sue Grand

Sue Grand presents a case of “a near-death clinical impasse,” conjuring “God at an impasse.” She questions the philosophical premises and culture biases that inform the foundations of psychoanalytic theory. She asks how we rewrite “the psychoanalytic subject.” This commentary explores the themes of clinical impasse, psychoanalysis and religion, martyrdom and self-sacrifice, the negative Oedipus or “Jacob complex,” multiplicity of selves, and psychoanalytic witnessing. Most important, it challenges the tendency to polarize the Jewish and Christian narratives such that Judaism is depicted as “this worldly” in contrast to Christianity, which is seen as celebrating martyrdom in identification with Christ. It argues for psychoanalysis to recognize the spiritual value of submission and surrender without splitting them into polar oppositions.     

Flashbacks,intrusions, mind-wandering – Instances of an involuntary memory spectrum: A commentary on Takarangi,Strange, and Lindsay (2014)

In their paper, Takarangi, Strange, and Lindsay (2014) showed in two experiments that participants who had witnessed a shocking film frequently “mind-wandered without awareness” about the content of the film. More importantly, they equated this effect with the occurrence of traumatic intrusions. In this commentary, we argue that the authors adhered to conceptually ambiguous terms, and thereby unintentionally contribute to an already existing conceptual blur in the trauma-memory field. We postulate that clear definitions are urgently needed for phenomena such as intrusions, flashbacks, and mind-wandering, when using them in the context of trauma memory. Furthermore, our proposal is that these phenomena can fall under a spectrum of different involuntary memory instances. We propose that by adopting stricter definitions and viewing them as separate, but interrelated phenomena, different lines of trauma-memory research can be reconciled, which would considerably advance the field.     

Walls and Holes in Psychosocial Research: From Psychoanalysis to Critique

This commentary reflects on the different innovative motifs introduced into psychosocial research by the contributions to this special issue: the risk of oversubjectification in research placing undue emphasis on the individual reasoning or feeling subject, the attempt to link the “feelings” and “talk” about emotion in one interpretative framework, the place of the interview in research which questions rather than reinforces “identity,” the location of subjects in a “place-assemblage” rather than in their own selves, and the reconfiguration of “mindfulness” so that it opens out to social relations rather than evades them. Focussing on the role of psychoanalysis in psychosocial research, I situate these motifs within the analysis of the machinery of “facialization” offered by Deleuze and Guattari, in which the “white wall” of signification is complemented and locked in place by the “black hole” of subjectivity.     

The visual influence of ostracism

Reflecting the fundamental human need to establish and maintain positive connections with others, it has been suggested that an Ostracism Detection System is sensitized to targets by which one has been ostracized. Evidence supporting the operation of this system has yet to be provided, however. Accordingly, using binocular rivalry to explore attentional processing, here, we considered the extent to which targets previously associated with ostracism dominate visual awareness. Participants initially performed a virtual ball‐tossing game (i.e., Cyberball) in which they were “ostracized” or “included” by the other players. Afterwards, the faces of these players were presented together with houses in a binocular rivalry task. The results revealed that targets associated with ostracism (vs. inclusion) dominated longest in visual awareness.     

Recognition memory and awareness: A large effect of study-test modalities on “know” responses following a highly perceptual orienting task

Abstract

Two experiments investigated the effects of varying the correspondence between presentation and test modes on states of conscious awareness in recognition memory. Experiment 1 used visual test items and showed that auditory and visual study modes differentially affected recollective experience and feelings of familiarity assessed by “remember” and “know” responses accompanying recognition: “know” responses were slightly enhanced following visual presentation, but this was largely offset by a similarly small increase in “remember” responses. Experiment 2 employed conditions designed to maximise any effect on “know” responses of the correspondence between presentation and test modes by using a highly perceptual orienting task at study. Under these conditions, there was a large mode correspondence effect on “know” responses. The results support the idea that “know” responses are particularly sensitive to perceptual factors.