Blind Field Research Articles

Detection of first- and second-order coherent motion in blindsight

Blindsight patients can detect fast moving stimuli presented within their blind field even when they deny any phenomenal visual experience. Although mounting evidence suggests the presence of different mechanisms and separate neural substrates underlying the processing of first-order (luminance-defined) and second-order (contrast-defined) motion, the perception of second-order motion in blindsight has scarcely been explored. In the present study, we investigated whether two blindsighted patients (GY and MS) can detect a variety of first- and second-order moving stimuli, and by using repetitive transcranial magnetic stimulation (rTMS), we assessed the role of V5/MT(+) and V3(+) in coherent motion processing. The hemianopes and four control subjects performed a two-interval forced-choice task in which they judged whether a pattern of coherently moving first-order or second-order textured squares moved in the first or second interval. They were not asked to report the direction of motion because neither of them could do so better than expected by chance. The results showed that MS, who has extensive destruction of the ventral cortical visual pathway as well as his V1 lesion, could not process second-order motion at all, whereas GY could perform second-order tasks but only at high-contrast modulation. This may have introduced first-order components in second-order moving stimuli and provided artifactual cues to motion. Moreover, rTMS delivered over area V5/MT(+) impaired detection of both first- and second-order motion in undamaged control subjects, whereas rTMS over V3(+) did not impair their performance in any of the stimuli employed. On the other hand, rTMS over V3(+) did impair GY's detection of first-order motion and high-contrast second-order moving textured squares that are likely to contain artifactual luminance cues. rTMS over V5/MT(+) impaired first-order motion detection in MS. Overall, the results suggest that neither of the blindsight patients can detect artifact-free second-order motion.

Awareness-related activity in prefrontal and parietal cortices in blindsight reflects more than superior visual performance

Many imaging studies report activity in the prefrontal and parietal cortices when subjects are aware as opposed to unaware of visual stimuli. One possibility is that this activity simply reflects higher signal strength or the superior task performance that is associated with awareness. To find out, we studied the hemianope GY who has unilateral destruction of almost all primary visual cortices. He exhibits 'blindsight', that is, he claims to have no conscious visual phenomenology (i.e., no visual qualia), for stationary stimuli presented to his right visual field (the blind field), although he can press keys to distinguish between different stimuli presented there. We presented to him a visual discrimination task, and equated performance for stimuli presented to the left or right visual field by presenting low contrast stimuli to his normal (left) field and high contrast stimuli to his blind (right) field. Superior accuracy can be a serious confound, and our paradigm allows us to control for it and avoid this confound. Even when performance was matched, and the signal strength was lower, visual stimulation to the normal (conscious) field led to higher activity in the prefrontal and parietal cortices. These results indicate that the activity in the prefrontal and parietal areas that has been reported in previous studies of awareness is not just due to a (signal strength or performance) confounds. One possibility is that it reflects the superior 'metacognitive' capacity that is associated with awareness, because GY was better able to distinguish between his own correct and incorrect responses for stimuli presented to his normal field than to his blind field.

Color experience in blindsight?

Blindsight, the ability to blindly discriminate wavelength and other aspects of stimuli in a blind field, sometimes occurs in people with lesions to striate (V1) cortex. There is currently no consensus on whether qualitative color information of the sort that is normally computed by double opponent cells in striate cortex is indeed computed in blindsight but doesn’t reach awareness, perhaps owing to abnormal neuron responsiveness in striate or extra-striate cortical areas, or is not computed at all. The existence of primesight, the experience of colored afterimages in blindsight, has been taken to suggest that qualitative color information is computed either in pre-striate or striate cortical areas but is not broadcast to working memory. I argue here that a recent study in which color phosphenes were induced in a blindsighter using bilateral transcranial magnetic stimulation indicates that computations necessary for conscious color vision are lost in blindsight. Owing to this loss, the neural responsiveness in extrastriate cortical areas is abnormal and hence is unable to give rise to color awareness. Blindsight is thus degraded vision in which the computations necessary for conscious color vision have been lost.

Hemianopic and Quadrantanopic Field Loss, Eye and Head Movements, and Driving

To compare eye and head movements, lane keeping, and vehicle control of drivers with hemianopic and quadrantanopic field defects with controls, and to identify differences in these parameters between hemianopic and quadrantanopic drivers rated safe to drive by a clinical driving rehabilitation specialist compared with those rated as unsafe. Eye and head movements and lane keeping were rated in 22 persons with homonymous hemianopic defects and 8 with quadrantanopic defects (mean age, 53 years) who were ≥6 months post-injury and 30 persons with normal fields (mean age, 53 years). All were licensed to drive and were current drivers or aimed to resume driving. Participants drove a 6.3-mile route along non-interstate city roads under in-traffic conditions. Vehicle control was assessed objectively by vehicle instrumentation for speed, braking, acceleration, and cornering. As a group, drivers with hemianopic or quadrantanopic defects drove slower, exhibited less excessive cornering or acceleration, and executed more shoulder movements than the controls. Those drivers with hemianopic or quadrantanopic defects rated as safe also made more head movements into their blind field, received superior ratings regarding eye movement extent and lane position stability, and exhibited less sudden braking and drove faster than those rated unsafe. Persons with hemianopic and quadrantanopic defects rated as safe to drive compensated by making more head movements into their blind field, combined with more stable lane keeping and less sudden braking. Future research should evaluate whether these characteristics could be trained in rehabilitation programs aimed at improving driving safety in this population.

Grasping the non-conscious: Preserved grip scaling to unseen objects for immediate but not delayed grasping following a unilateral lesion to primary visual cortex

Patients with damage to primary visual cortex can sometimes direct actions towards ‘unseen’ targets located in areas of the visual field that are deemed ‘blind’ on the basis of static perimetry tests. Here, we show that a patient with a complete right homonymous hemianopia after a V1 lesion remains sensitive to the width of objects presented in her blind field but only when reaching out to grasp them in ‘real-time’. A subsequent fMRI experiment revealed spared extra-geniculostriate pathways, which may mediate her preserved abilities. Taken together, the results support the view that visually guided movements can be mediated by pathways that do not support visual consciousness.

Task-Irrelevant Blindsight and the Impact of Invisible Stimuli

Despite their subjective invisibility, stimuli presented within regions of absolute cortical blindness can both guide forced-choice behavior when they are task-relevant and modulate responses to visible targets when they are task-irrelevant. We here tested three hemianopic patients to learn whether their performance in an attention-demanding rapid serial visual presentation task would be affected by task-irrelevant stimuli. Per trial, nine black letters and one white target letter appeared briefly at fixation; the white letter was to be named at the end of each trial. On 50% of trials, a task-irrelevant disk (−0.6 log contrast) was presented to the blind field; in separate blocks, the same or a very low negative contrast distractor was presented to the sighted field. Mean error rates were high and independent of distractor condition, although the high-contrast sighted-field disk impaired performance significantly in one participant. However, when trials with and without distractors were considered separately, performance was most impaired by the high-contrast disk in the blind field, whereas the same disk in the sighted field had no effect. As this disk was least visible in the blind and most visible in the sighted field, attentional suppression was inversely related to visibility. We suggest that visual awareness, or the processes that generate it and are compromised in the blind hemisphere, enhances or enables effective attentional suppression.

Saccadic Remapping in Hemianopia

We experience visual stability despite shifts of the visual array across the retina produced by eye movements. The visual system predicts the retinal consequences of saccades through a process known as saccadic remapping. We explored remapping in damaged visual cortex by presenting a stimulus in the blind field of a patient with hemianopia. Reported awareness was elevated when a saccade was about to bring the stimulated location into the sighted field, even though the stimulus was removed before the saccade began and so never stimulated the sighted field. The presence of a blind-field stimulus before saccade also increased sensitivity to a near-threshold sighted-field stimulus that was presented at the same spatial location after saccade. The results suggest early visual areas are not necessary for remapping, but play a role in verifying predictions about the retinal positions of post-saccadic targets generated by remapping.

Illusory motion perception in blindsight

Motion detection is typically spared in blindsight, which results from damage to the striate cortex (area V1) of the brain that is sufficient to eliminate conscious visual awareness and severely reduce sensitivity to luminance contrast, especially for high spatial and low temporal frequencies. Here we show that the discrimination of motion direction within cortically blind fields is not attributable to feature tracking (the detection of changes in position or shape), but is due instead to the detection of first-order motion energy (spatiotemporal changes in luminance). The key to this finding was a version of the line motion illusion entailing reverse-phi motion in which opposing motion directions are simultaneously cued by motion energy and changes in stimulus shape. In forced-choice tests, a blindsighted test subject selected the direction cued by shape change when the stimulus was presented in his intact field, but reliably selected the direction cued by motion energy when the same stimulus was presented in his blind field, where relevant position information was either inaccessible or invalid. Motion energy has been characterized as objectless, so reliance on motion energy detection is consistent with impaired access to shape information in blindsight. The dissociation of motion direction by visual field (cortically blind vs. intact) provides evidence that two pathways from the retina to MT/V5 (the cortical area specialized for motion perception) are functionally distinct: the retinogeniculate pathway through V1 is specialized for feature-based motion perception, whereas the retinocollicular pathway, which bypasses V1, is specialized for detecting motion energy.

The primary visual cortex, and feedback to it, are not necessary for conscious vision

A compelling single case report of visual awareness (visual qualia) without primary visual cortex would be sufficient to refute the hypothesis that the primary visual cortex and the back-projections to it are necessary for conscious visual experience. In a previous study, we emphasized the presence of crude visual awareness in Patient G.Y., with a lesion of the primary visual cortex, who is aware of, and able to discriminate, fast-moving visual stimuli presented to his blind field. The visual nature of Patient G.Y.'s blind field experience has since been questioned and it has been suggested that the special circumstances of repeated testing over decades may have altered Patient G.Y.'s visual pathways. We therefore sought new evidence of visual awareness without primary visual cortex in patients for whom such considerations do not apply. Three patients with hemianopic field defects (Patient G.N. and Patient F.B. with MRI confirmed primary visual cortex lesions, Patient C.G. with an inferred lesion) underwent detailed psychophysical testing in their blind fields. Visual stimuli were presented at different velocities and contrasts in two- and four-direction discrimination experiments and the direction of motion and awareness reported using a forced-choice paradigm. Detailed verbal reports were also obtained of the nature of the blind field experience with comparison of the drawings of the stimulus presented in the blind and intact fields, where possible. All three patients reported visual awareness in their blind fields. Visual awareness was significantly more likely when a moving stimulus was present compared to no stimulus catch trials (P < 0.01 for each subject). Psychophysical performance in Patient F.B. and Patient G.N. was consistent with the Riddoch syndrome, with higher levels of visual awareness for moving compared to static stimuli (P < 0.001) and intact direction discrimination (P < 0.0001 for two- and four-direction experiments). Although the blind field experience of all three subjects was degraded, it was clearly visual in nature. We conclude that the primary visual cortex or back-projections to it are not necessary for visual awareness.

Rapid Adaptation of Visual Search in Simulated Hemianopia

Patients with homonymous hemianopia have altered visual search patterns, but it is unclear how rapidly this develops and whether it reflects a strategic adaptation to altered perception or plastic changes to tissue damage. To study the temporal dynamics of adaptation alone, we used a gaze-contingent display to simulate left or right hemianopia in 10 healthy individuals as they performed 25 visual search trials. Visual search was slower and less accurate in hemianopic than in full-field viewing. With full-field viewing, there were improvements in search speed, fixation density, and number of fixations over the first 9 trials, then stable performance. With hemianopic viewing, there was a rapid shift of fixation into the blind field over the first 5-7 trials, followed by continuing gradual improvements in completion time, number of fixations, and fixation density over all 25 trials. We conclude that in the first minutes after onset of hemianopia, there is a biphasic pattern of adaptation to altered perception: an early rapid qualitative change that shifts visual search into the blind side, followed by more gradual gains in the efficiency of using this new strategy, a pattern that has parallels in other studies of motor learning.

Consciousness of the first order in blindsight

At suprathreshold levels, detection and awareness of visual stimuli are typically synonymous in nonclinical populations. But following postgeniculate lesions, some patients may perform above chance in forced-choice detection paradigms, while reporting not to see the visual events presented within their blind field. This phenomenon, termed "blindsight," is intriguing because it demonstrates a dissociation between detection and perception. It is possible, however, for a blindsight patient to have some "feeling" of the occurrence of an event without seeing per se. This is termed blindsight type II to distinguish it from the type I, defined as discrimination capability in the total absence of any acknowledged awareness. Here we report on a well-studied patient, D.B., whose blindsight capabilities have been previously documented. We have found that D.B. is capable of detecting visual patterns defined by changes in luminance (first-order gratings) and those defined by contrast modulation of textured patterns (textured gratings; second-order stimuli) while being aware of the former but reporting no awareness of the latter. We have systematically investigated the parameters that could lead to visual awareness of the patterns and show that mechanisms underlying the subjective reports of visual awareness rely primarily on low spatial frequency, first-order spatial components of the image.

Plasticity of the visual system after early brain damage

The aim of this review is to discuss the existing evidence supporting different processes of visual brain plasticity after early damage, as opposed to damage that occurs during adulthood. There is initial evidence that some of the neuroplastic mechanisms adopted by the brain after early damage to the visual system are unavailable at a later stage. These are, for example, the ability to differentiate functional tissue within a larger dysplastic cortex during its formation, or to develop new thalamo-cortical connections able to bypass the lesion and reach their cortical destination in the occipital cortex. The young brain also uses the same mechanisms available at later stages of development but in a more efficient way. For example, in people with visual field defects of central origin, the anatomical expansion of the extrastriatal visual network is greater after an early lesion than after a later one, which results in more efficient mechanisms of visual exploration of the blind field. A similar mechanism is likely to support some of the differences found in people with blindsight, the phenomenon of unconscious visual perception in the blind field. In particular, compared with people with late lesions, those with early brain damage appear to have stronger subjective awareness of stimuli hitting the blind visual field, reported as a conscious feeling that something is present in the visual field. Expanding our knowledge of these mechanisms could help the development of early therapeutic interventions aimed at supporting and enhancing visual reorganization at a time of greatest potential brain plasticity.

Visual field expansion after visual restoration therapy

Objective: To determine whether visual field expansion occurs with visual restoration therapy (VRT), using fundus-controlled microperimetry to assess visual fields. Design: This longitudinal cohort analysis assesses patients’ visual fields before and after visual restoration therapy using microperimetry and standard high-resolution perimetry. Subjects: Seven patients with stroke-induced homonymous field cuts were studied. Intervention: Visual restoration therapy is a computerized, home-based treatment aimed at reducing the size of the visual field defect of stroke patients with hemianopia through repetitive stimulation of the visual borderzone adjacent to the blind field. During twice-daily therapy for three months patients maintain central fixation while responding to eccentrically placed stimuli in the visual borderzone. The programme is adjusted monthly to changes in the patient’s visual field. Controversy exists as to whether expansion of visual fields measured at home with high-resolution perimetry is due to inadvertent eye movements and therefore would overrepresent the treatment’s effect. Main measures: Microperimentry uses an infrared camera to track retinal vessels so that any shift or movement between the reference image and the real-time fundus image corrects the stimulus position, thus delivering stimuli to known retinal locations, and allowing accurate assessment of visual fields independent of eye movements. Results: There was an average improvement in stimulus detection rate by microperimetry of 12.5% (range —1.4% to 38.9%, P =0.033). Six of 7 patients had ≥ 3% improvement in stimulus detection by home-based perimetry. Conclusion: Our results demonstrate modest but real expansion in visual fields following visual restoration therapy which is not due to eye movements.

Improved detection following Neuro-Eye Therapy in patients with post-geniculate brain damage

Damage to the optic radiation or the occipital cortex results in loss of vision in the contralateral visual field, termed partial cortical blindness or hemianopia. Previously, we have demonstrated that stimulation in the field defect using visual stimuli with optimal properties for blindsight detection can lead to increases in visual sensitivity within the blind field of a group of patients. The present study was aimed to extend the previous work by investigating the effect of positive feedback on recovery of visual sensitivity. Patients' abilities for detection of a range of spatial frequencies within their field defect were determined using a temporal two-alternative forced-choice technique, before and after a period of visual training (n=4). Patients underwent Neuro-Eye Therapy which involved detection of temporally modulated spatial grating patches at specific retinal locations within their field defect. Three patients showed improved detection ability following visual training. Based on our previous studies, we had hypothesised that should the occipital brain lesion extend anteriorly to the thalamus, little recovery would be expected. Here, we describe one such case who showed no improvements after extensive training. The present study provides further evidence that recovery (a) can be gradual and may require a large number of training sessions (b) can be accelerated using positive feedback and (c) may be less likely to take place if the occipital damage extends anteriorly to the thalamus.

Blindsight and enumeration: A case study

“Blindsight” is a term first coined by Weiskrantz et al. in 1974 to describe residual visual performance in the cortically blind. It has been postulated that blindsight could be due to the retinotectal pathway projecting information past V1 to later cortical structures. Our interest was in whether the pathways responsible for blindsight could also support enumeration. We tested a 53-year-old male, C.H., who had suffered a medial right occipital lobe stroke seven months previous. The stroke resulted in an upper left homonymous quadrantanopia. In order to determine whether there were any residual abilities in the blind field, we first tested basic detection and discrimination skills. C.H. was able to determine whether or not a 7° X 7° visual angle object was presented in his blind field with near perfect accuracy though his accuracy was only 90% for smaller (4.6° X 2.2°) figures. C.H. also discriminated between large X′s and O′s and horizontal and vertical bars with good accuracy when the figures were large (78% for X vs. O and 73% for horizontal vs. vertical bars). Throughout these tests C.H. staunchly maintained that he could not see the stimuli and that he was only guessing. Because it was clear that C.H. had some residual ability in his blind field, we tested his enumeration. The enumeration task involved 1-3 items. These items were 1.5° Blindsight and enumeration: A case study http://www.journalofvision.org/content/10/7/898.short 1 of 2 2015-01-03, 8:42 PM X 1.5° black diamonds, 0-2 in his blind field and 1-3 in his non-blind field. Across all conditions, C.H. was able to use the information in his blind field and identify the total number of items presented at levels of performance that were well above chance. Because C.H. appears to use blind field information to enumerate, we postulate that enumeration ability may be mediated by the same structures that support blindsight. Received June 26, 2010. © 2010 ARVO Articles citing this article Blliindsiight and enumeratiion:: A case study J Vis August 12, 2010 10(7): 898 Abstract Blindsight and enumeration: A case study http://www.journalofvision.org/content/10/7/898.short 2 of 2 2015-01-03, 8:42 PM

Anomalous global effects induced by ‘blind’ distractors in visual hemifield defects

Previous research has revealed that a stimulus presented in the blind visual field of participants with visual hemifield defects can evoke oculomotor competition, in the absence of awareness. Here we studied three cases to determine whether a distractor in a blind hemifield would be capable of inducing a global effect, a shift of saccade endpoint when target and distractor are close to each other, in participants with lesions of the optic radiations or striate cortex. We found that blind field distractors significantly shifted saccadic endpoints in two of three participants with lesions of either the striate cortex or distal optic radiations. The direction of the effect was paradoxical, however, in that saccadic endpoints shifted away from blind field distractors, whereas endpoints shifted towards distractors in the visible hemifields, which is the normal global effect. These results provide further evidence that elements presented in the blind visual field can generate modulatory interactions in the oculomotor system, which may differ from interactions in normal vision.

Line bisection in homonymous visual field defects – Recent findings and future directions

Homonymous visual field defects are a frequent consequence of brain damage occurring in some 20–40% of stroke patients. They are often accompanied by a peculiar spatial bias termed “Hemianopic Line Bisection Error” (HLBE). Although known for more than 100 years the explanations for the HLBE put forward remain controversial. One explanation holds that the HLBE is a direct consequence of the field defect itself and reflects a compensatory shift of attention towards the scotoma. Another, contradicting position states that although the HLBE is frequently found in any type of homonymous visual field defect – not only hemianopia – it is not simply a direct consequence of the field defect itself, although it does contribute to it. According to this position, the HLBE arises from additional damage to extrastriate cortex, thus causing the spatial bias towards the blind field. In the present article we summarize the main arguments of both theoretical positions and argue that although both accounts are valid, they are incomplete and several important issues remain unresolved. These include the potential contribution of eccentric fixation to the HLBE, the question of multimodal impairments, the role of (visuo)-motor processes, the relation between the HLBE and visual field recovery, and the exact clinical significance of the HLBE. Thus, far from concluding the research on the line-bisection error in hemianopia, the recent series of publications on this topic serve as a welcome reminder of how much more research is needed.

Reflections on blindsight: Neuroimaging and behavioural explorations clarify a case of reversed localisation in the blind field of a patient with hemianopia.

Blindsight refers to residual visual abilities of patients with primary visual cortex lesions. Most of this research uses single case studies, most famously patient GY. We examined a patient (DC) after surgical resection of V1 who demonstrated robust but reversed blind field target localisation, mislocalising midline blind field targets to the periphery and vice versa. This pattern was reliable across multiple sessions and was not because of extraocular light scatter. We then used functional magnetic resonance imaging to examine neural responses to blind field motion stimuli with no evidence of motion-selective activation in DC's extrastriate cortex in the damaged hemisphere, in stark contrast to GY who showed robust bilateral activation in response to blind field stimuli. This suggests that DC's blind field performance may not represent true blindsight. Follow-up testing with the target--background contrast reversed (i.e., black targets/white background), eliminated DC's reversed localisation, strongly suggesting that she was employing an unusual decision criterion based on intraocular light scatter. DC's failure to demonstrate true blindsight may be related to the age at which she acquired her lesion--much later in life than GY.

Collicular Vision Guides Nonconscious Behavior

Following destruction or deafferentation of primary visual cortex (area V1, striate cortex), clinical blindness ensues, but residual visual functions may, nevertheless, persist without perceptual consciousness (a condition termed blindsight). The study of patients with such lesions thus offers a unique opportunity to investigate what visual capacities are mediated by the extrastriate pathways that bypass V1. Here we provide evidence for a crucial role of the collicular-extrastriate pathway in nonconscious visuomotor integration by showing that, in the absence of V1, the superior colliculus (SC) is essential to translate visual signals that cannot be consciously perceived into motor outputs. We found that a gray stimulus presented in the blind field of a patient with unilateral V1 loss, although not consciously seen, can influence his behavioral and pupillary responses to consciously perceived stimuli in the intact field (implicit bilateral summation). Notably, this effect was accompanied by selective activations in the SC and in occipito-temporal extrastriate areas. However, when instead of gray stimuli we presented purple stimuli, which predominantly draw on S-cones and are thus invisible to the SC, any evidence of implicit visuomotor integration disappeared and activations in the SC dropped significantly. The present findings show that the SC acts as an interface between sensory and motor processing in the human brain, thereby providing a contribution to visually guided behavior that may remain functionally and anatomically segregated from the geniculo-striate pathway and entirely outside conscious visual experience.

Blindsight Mediated by an S-Cone-independent Collicular Pathway: An fMRI Study in Hemispherectomized Subjects

The purpose of our study was to investigate the ability to process achromatic and short-wavelength-sensitive cone (S-cone)-isolating (blue-yellow) stimuli in the blind visual field of hemispherectomized subjects and to demonstrate that blindsight is mediated by a collicular pathway that is independent of S-cone inputs. Blindsight has been described as the ability to respond to visual stimuli in the blind visual field without conscious awareness [Weiskrantz, L., Warrington, E. K., Sanders, M. D., & Marshall, J. Visual capacity in the hemianopic field following a restricted occipital ablation. Brain, 97, 709-728, 1974]. The roles of the subcortical neural structures in blindsight, such as the pulvinar and the superior colliculus, have been debated and an underlying neural correlate has yet to be confirmed. Using fMRI, we tested the ability to process visual stimuli that isolated the achromatic and short-wavelength-sensitive (S-)-cone pathways in three subjects: one control subject, one hemispherectomized subject with blindsight, and one hemispherectomized subject without blindsight. We demonstrated that (1) achromatic and S-cone-isolating stimuli presented to the normal visual hemifield of hemispherectomized subjects and to both visual hemifields of the control subject activated contralateral visual areas (V1/V2), as expected; (2) achromatic stimulus presentation but not S-cone-isolating stimulus presentation to the blind hemifield of the subject with blindsight activated visual areas FEF/V5; (3) whereas the cortical activation of the control subject was enhanced by an additional stimulus (achromatic and S-cone isolating) presented in the contralateral visual field, activation pattern of the subject with blindsight was enhanced by achromatic stimuli only. We conclude that the human superior colliculus is blind to the S-cone-isolating stimuli, and blindsight is mediated by an S-cone-independent collicular pathway.