Variability of residual vision in hemianopic subjects

Using stabilized visual field mapping techniques, seven hemianopic subjects were extensively investigated for residual visual abilities. Isolated islands of detection abilities were demonstrated by four of these subjects. Additional abilities demonstrated within these islands included saccadic and verbal localization, wavelength discrimination, form discrimination, and motion detection. These abilities were also accompanied by low-confidence ratings, and thus have the character of blindsight. It is noteworthy that different subjects demonstrated different abilities at different visual field locations, underscoring the between and within subject variability often observed with blindsight. Furthermore, magnetic resonance images obtained for each subject demonstrated variable sparing of occipital cortex. Such cortical sparing, in conjunction with the behavioral variability, supports the notion that some instances of blindsight are mediated by remnants of the primary visual pathway.     

Neural activity within area V1 reflects unconscious visual performance in a case of blindsight

Although lesions of the striate (V1) cortex disrupt conscious vision, patients can demonstrate surprising residual abilities within their affected visual field, a phenomenon termed blindsight. The relative contribution of spared "islands" of functioning striate cortex to residual vision, versus subcortical pathways to extrastriate areas, has implications for the role of early visual areas in visual awareness and performance. Here we describe the behavioral and neural features of residual cortical function in Patient M.C., who sustained a posterior cerebral artery stroke at the age of 15 years. Within her impaired visual field, we found preserved visual abilities characteristic of blindsight, including superior detection of motion, and above-chance discrimination of shape, color, and motion direction. Functional magnetic resonance imaging demonstrated a retinotopically organized representation of M.C.'s blind visual field within the lesioned occipital lobe, specifically within area V1. The incongruity of a well-organized cortex and M.C.'s markedly impaired vision was resolved by measurement of functional responses within her damaged occipital lobe. Attenuated neural contrast-response functions were found to correlate with M.C.'s impaired psychophysical performance. These results demonstrate that the behavioral features of blindsight may arise in the presence of residual striate responses that are spatially organized and sensitive to contrast variation.     

Absence of S-cone input in human blindsight following hemispherectomy

Destruction of the occipital cortex presumably leads to permanent blindness in the contralateral visual field. Residual abilities to respond to visual stimuli in the blind field without consciously experiencing them have, however, been described in cortically blind patients and are termed 'blindsight'. Although the neuronal basis of blindsight remains unknown, possible neuronal correlates have been proposed based on the nature of the residual vision observed. The most prominent but still controversial hypothesis postulates the involvement of the superior colliculi in blindsight. Here we demonstrate, using a computer-based reaction time test in a group of hemispherectomized subjects, that human 'attention-blindsight' can be measured for achromatic stimuli but disappears for stimuli that solely activate S-cones. Given that primate data have shown that the superior colliculi lacks input from S-cones, our results lend strong support to the hypothesis that 'attention-blindsight' is mediated through a collicular pathway. The contribution of a direct geniculo-extrastriate-koniocellular projection was ruled out by testing hemispherectomized subjects in whom a whole hemisphere has been removed or disconnected for the treatment of epilepsy. A direct retino-pulvinar-cortical connection is also unlikely as the pulvinar nucleus is known to receive input from S-cones as well as from L/M-cone-driven colour-opponent ganglion cells.     

A temporal/nasal asymmetry for blindsight in a localisation task: evidence for extrageniculate mediation

Some patients with hemianopia due to striate cortex lesions show above chance ability in reporting visual stimuli presented in the blind visual field, a phenomenon commonly known as blindsight. Here we report a patient with a dense right hemianopia whose blindsight shows a temporal/nasal asymmetry. MP was tested in a two-alternative forced-choice localisation task, with either the right eye or the left eye patched in separate blocks. When targets appeared in the contralesional temporal hemifield, MP's localisation performance was extremely accurate, whilst she performed at chance with targets in the contralesional nasal hemifield. This is the first demonstration of a temporal/nasal asymmetry for blindsight in a forced-choice paradigm, and is consistent with blindsight in MP's hemianopic field being mediated by a subcortical, extrageniculate route.     

Blindsight after optic nerve injury indicates functionality of spared fibers

Some patients with lesions in the geniculostriate pathway (GSP) can respond to visual stimuli in the blind field without conscious acknowledgement. The substrate for this "blindsight" is controversial: whether it is the uninjured extrastriate pathway (EXP), which bypasses the lesion site, or residual fibers within damaged visual cortex ("islands of vision"). Using stimulus detection, localization, and spatial summation tasks, we have found blindsight in patients with damage both in the optic nerve (ON) and EXP. The prevalence and functional characteristics of their blindsight are indistinguishable from that in patients with GSP lesions, so blindsight does not require a completely intact EXP. The present findings support the view that a few surviving ON axons within an area of primary damage are sufficient to mediate blindsight: Several combinations of partially intact pathways can transmit information to the extrastriate cortex and the sum of activation of all visual fibers surviving the injury determines if and to what extent blindsight occurs.     

Making the blindsighted see

A lesion of striate cortex, area V1, produces blindness in the retinotopically corresponding part of the visual field, although in some cases visual abilities in the blind field remain that are paradoxically devoid of conscious visual percepts (“blindsight”). Here we demonstrate that the blindsight subject GY can experience visual sensations of phosphenes in his blind field induced by transcranial magnetic stimulation (TMS). Such blind field percepts could only be induced when stimulation was applied bilaterally, i.e. over GY's area V5/MT in both hemispheres. Consistent with an earlier report [Cowey, A., & Walsh, V. (2000). Magnetically induced phosphenes in sighted, blind and blindsighted observers. Neuroreport, 11, 3269–3273], GY never experienced phosphenes when stimulation was restricted to his ipsilesional V5/MT. To the best of our knowledge this is the first time GY has experienced visual qualia in his blind hemifield. The present report characterizes the necessary conditions for such conscious experience in his hemianopic visual field and interprets them as demonstrating that only via a contribution from GY's intact hemisphere can activation in the damaged hemisphere reach visual awareness.     

Visual attention in blindsight: sensitivity in the blind field increased by targets in the sighted field

Damage to primary visual cortex results in a scotoma, or region of blindness, which can nevertheless be accompanied by residual visual abilities in the affected field. This dissociation of visual processing and visual experience is often described as blindsight. We investigated the role of visual attention in blindsight and the application of limited visual resources between the blind and sighted fields in a well-studied case. Contrary to expectations based on normal capacity limits, processing a target in the sighted field improved rather than interfered with processing of targets in the blind field. The benefit from the sighted field was greater when blind and sighted targets were in symmetric locations about the vertical meridian.     

Spatial and temporal processing in a subject with cortical blindness following occipital surgery

Blindsight subjects are typically better at discriminating rapid, transient visual events than those with gradual on/off-sets. Surprisingly, the detailed investigation of temporal characteristics of mechanisms mediating blindsight is only reported in one subject (GY). It is of interest to establish whether these characteristics are similar to those in other cases of blindsight. Here, we report on a systematic study of spatio-temporal properties of mechanisms mediating blindsight in a subject VN. VN has a lower right quadranopia following surgical removal of the left occipital cortex above the calcarine sulcus, therefore, there are no remaining islands of intact visual cortex within this area. Similar to GY, the blindsight mechanisms in VN have narrowly tuned band-pass temporal characteristics with a peak sensitivity at 20Hz and above chance performance at temporal frequencies >/=10 and 相似文献   

Visual restoration in cortical blindness: insights from natural and TMS-induced blindsight

Unilateral damage to visual cortex of the parietal or occipital lobe can cause the patient to be unaware of contralesional visual information due to either hemispatial neglect or hemianopia. It is now known that both neglect and hemianopia result from the disruption of a dynamic interaction between cortical visual pathways and more phylogenetically primitive visual pathways to the midbrain. We consider the therapeutic implications of these cortical-subcortical interactions in the rehabilitation of hemianopia. We start with the pheonmenon of "blindsight", in which patients with hemianopia can be shown, by implicit measures of visual detection or discrimination, to process visual information without conscious awareness. Some variants of blindsight have been postulated to recruit subcortical processes, while others may reflect compensatory optimisation of processing of spared visual cortex. Both mechanisms may offer opportunities for innovative strategies for rehabilitation of visual field defects. We relate the neural mechanisms that have been proposed to underlie blindsight to those that have been suggested to underlie the recovery of visual function after rehabilitation. It is suggested that the similarity and overlap of the neural processes supporting blindsight and recovery of visual function might provide insights for effective rehabilitation strategies for restoring visual functions.     

Pathological perceptual completion in hemianopia extends to the control of reach-to-grasp movements

The neuropsychological phenomenon of blindsight is observed when patients who are cortically blind exhibit residual visual processing capabilities for stimuli presented within their scotoma to which they are otherwise unaware. Cortically blind patients may also exhibit the phenomenon of pathological visual completion in which, paradoxically, they can become aware of a complete visual stimulus even when a significant portion of that stimulus falls within their blind hemifield. In this study, the ability of a blindsight patient (G.Y.) to use visual information to control reach-to-grasp movements to static objects presented within his blind hemifield was investigated. The results indicate that while G.Y. was insensitive to variations in object size when reaching for objects presented entirely within his blind hemifield, his ability to accurately grasp objects located within his blind field was vastly improved if part of the object to be grasped extended into his seeing hemifield. This finding demonstrates that visual awareness can facilitate the visuomotor processing of object form within G.Y.'s apparently blind field, and suggests that the primary deficit in blindsight may be an impairment of visual consciousness rather than an absolute loss of visual function.     

Defensive responses to looming visual stimuli in monkeys with unilateral striate cortex ablation.

A number of residual visual functions including detection, localization and discrimination of visual stimuli, have been demonstrated in the "blind" fields of monkeys and human patients following damage to striate cortex. We report here that avoidance movements of the head can also be elicited from monkeys with unilateral striate cortex ablations when a "looming" stimulus is presented within the hemianopic and presumably "blind" field. The possible role of the retinofugal projection to superior colliculus in the mediation of these defensive head movements is discussed.     

Residual vision in a subject with damaged visual cortex.

It is well known that a lesion in the optic radiation or striate cortex leads to blind visual regions in the retinotopically corresponding portion of the visual field. However, various studies show that some subjects still perceive certain stimuli even when presented in the "blind" visual field. Such subjects either perceive stimuli abnormally or only certain aspects of them (residual vision) or, in some cases, deny perception altogether even though visual performance can be shown to be above chance (blindsight). Research on monkeys has suggested a variety of parallel extrastriate visual pathways that could bypass the striate cortex and mediate residual vision or blindsight. In the present study, we investigated a subject with perimetrically blind visual areas caused by bilateral brain damage. Black and white stimuli were presented at many locations in the intact and affected areas of the visual field. The subject's task was to state, using confidence levels, whether the target stimulus was black or white. The results revealed an area in the "blind" visual field in which the subject perceived a light flash when the experimental black stimulus was presented. We hypothesize that a spared region in the visual cortex most likely accounts for these findings.     

Blindsight in action: what can the different sub-types of blindsight tell us about the control of visually guided actions?

Blindsight broadly refers to the paradoxical neurological condition where patients with a visual field defect due to a cortical lesion nevertheless demonstrate implicit residual visual sensitivity within their field cut. The aim of this paper is twofold. First, through a selective review of the blindsight literature we propose a new taxonomy for the subtypes of residual abilities described in blindsight. Those patients able to accurately act upon blind field stimuli (e.g. by pointing or saccading towards them) are classified as having 'action-blindsight', those whose residual functions can be said to rely to some extent upon attentive processing of blind field stimuli are classified as demonstrating 'attention-blindsight', while finally, patients who have somewhat accurate perceptual judgements for blind field stimuli despite a complete lack of any conscious percept, are classified as having 'agnosopsia'--literally meaning 'not knowing what one sees'. We also address the possible neurological substrates of these residual sensory processes. Our second aim was to investigate the most striking subtype of blindsight, action-blindsight. We review the data relevant to this subtype and the hypotheses proposed to account for it, before speculating on how action-blindsight may inform our normal models of visuomotor control.     

Spectral sensitivity in hemianopic macaque monkeys.

We measured the increment threshold sensitivity to 2 degrees, 200-ms targets presented at a lateral and radial eccentricity of approximately 20-26 degrees in both visual hemifields of three macaque monkeys whose left striate cortex had been removed 5 years earlier, and in one normal control. As in patients with blindsight, sensitivity of the hemianopic field for blue, green and red stimuli was reduced by as little as 0.5 log units. With increasing light adaptation from scotopic to mesopic to photopic levels, there was a progressive increase in the sensitivity to long wavelengths relative to that for short and medium wavelengths. This shift in relative sensitivity ('Purkinje shift') shows that rod and cone mechanisms operate in both the normal and hemianopic fields and that the sensitivity that remains following removal of striate cortex is not mediated exclusively by rods.     

The neurobiology of blindsight.

Some patients can respond to visual stimuli presented within their clinically absolute visual field defects that have been caused by partial destruction of striate cortex. This puzzling phenomenon of looking, pointing, detecting and discriminating without seeing has been called blindsight, and has fascinated philosophers and neuroscientists alike as a spotlight on the nature of unconscious or covert awareness, and the means it provides of studying the visual information carried by pathways other than the major route through the striate cortex.     

The role of spared calcarine cortex and lateral occipital cortex in the responses of human hemianopes to visual motion

Some patients, who are rendered perimetrically blind in one hemifield by cortical lesions, nevertheless exhibit residual visual capacities within their field defects. The neural mechanism that mediates the residual visual responses has remained the topic of considerable debate. One explanation posits the subcortical visual pathways that bypass the primary visual cortex and innervate the extrastriate visual areas as the substrate that underlies the residual vision. The other explanation is that small islands of the primary visual cortex remain intact and provide the signals for residual vision. We have performed behavioral and functional magnetic resonance imaging experiments to investigate the validity of the two explanations of residual vision. Our behavioral experiments indicated that of the seven hemianopes tested, two had the ability to discriminate the direction of a drifting grating. This residual visual response was shown with fMRI to be the result of spared islands of calcarine cortical activity in one of the hemianopes, whereas only lateral occipital activity was documented in the other patient. These results indicate that the underlying neural correlates of residual vision can vary between patients. Moreover, our study emphasizes the necessity of ruling out the presence of islands of preserved function and primary visual cortex before assigning residual visual capacities to the properties of visual pathways that bypass the primary visual cortex.     

Spatial channels of visual processing in cortical blindness

Blindsight is the ability of some cortically blind patients to discriminate visual events presented within their field defect. We have examined a fundamental aspect of visual processing, namely the detection of spatial structures presented within the field defect of 10 cortically blind patients. The method outlined is based on the detection of high-contrast stimuli and is effective in flagging a 'window of detection' in the spatial frequency spectrum, should it exist. Here we report on the presence of a narrowly tuned psychophysical spatial channel optimally responding to frequencies less than 4 cycles/ degrees in eight out of 10 patients tested. The two patients who did not show any evidence of blindsight appear to have intact midbrain structures, but have lesions that extend from the occipital cortex to the thalamus. In addition, we have recorded subjective reports of awareness of the visual events in each trial. Detection scores of eight blindsight patients were subsequently subdivided based on the subjective reports of awareness. It appears that the psychophysical spatial channel-mediating responses in the absence of any awareness of the visual event have a narrower frequency response than those involved when the patients report some awareness of the visual event. The findings are discussed in relation to previous reports on the incidence of blindsight and performance on tasks involving spatial processing.     

A blindsight conundrum: how to respond when there is no correct response

Whereas research on blindsight customarily defines the correct responses to all visual stimuli presented to the cortically blind field, we here introduced a small number of unexpected 'no stimulus' trials in a localization task, to discover whether they would elicit the same responses as blind field targets. As no correct responses existed for the blank stimuli, our subjects, three hemianopic and one normal monkey, and one human hemianope who was aware of many blind-field targets, could either respond to these catch trials as to a target or refrain from responding. Visual stimuli were presented singly at four possible positions, two in the blind field of the hemianopes, and all subjects correctly localized the vast majority of targets in either hemifield. On blank trials, the monkeys, but not the human, often failed to respond, and when they did respond, all hemianopes almost invariably touched a target position in the blind field. Analysis of reaction times showed that necessarily false responses to blank stimuli took longer than responses to blind field targets. However, apart from one hemianopic monkey, incorrect target responses took as long as responses to blank stimuli. The human hemianope showed the same pattern of reaction times as the hemianopic monkeys unless he had to report on stimulus awareness and confidence. Then, his confidence reports and response times mirrored his awareness of the stimuli, but neither differed for correct versus false responses once these were separated for 'aware' versus 'unaware' trials. The hemianopic monkeys' response probability and reaction time data indicate that they, implicitly or explicitly, registered differences between target and blank stimuli and, in one case, even between false responses to blind-field and blank stimuli.     

Non-conscious recognition of affect in the absence of striate cortex

Functional neuroimaging experiments have shown that recognition of emotional expressions does not depend on awareness of visual stimuli and that unseen fear stimuli can activate the amygdala via a colliculopulvinar pathway. Perception of emotional expressions in the absence of awareness in normal subjects has some similarities with the unconscious recognition of visual stimuli which is well documented in patients with striate cortex lesions (blindsight). Presumably in these patients residual vision engages alternative extra-striate routes such as the superior colliculus and pulvinar. Against this background, we conjectured that a blindsight subject (GY) might recognize facial expressions presented in his blind field. The present study now provides direct evidence for this claim.     

Unlike in Clinical Blindsight Patients,Unconscious Processing of Chromatic Information Depends on Early Visual Cortex in Healthy Humans

BackgroundLarge amount of data concerning the neural mechanisms that mediate unconscious vision come from cortically blind patients who can process stimuli presented in their blind visual field. How well the findings generalize to neurologically healthy humans remains open.ObjectiveUsing transcranial magnetic stimulation (TMS), we studied whether chromatic processing depends on the early visual cortex in healthy participants.MethodWe employed a phenomenon called the redundant target effect (RTE): simple reaction times to two stimuli are faster than to one stimulus, even when one of the stimuli is presented outside consciousness.ResultsThe RTE produced by chromatically defined stimuli disappeared when the contralateral stimulus of two bilateral stimuli was suppressed from consciousness.ConclusionsIn contrast to studies on blindsight patients, the results imply that the early visual cortex is necessary for the processing of chromatic information in neurologically healthy humans.