Blindsight and visual awareness

Some patients with damaged striate cortex have blindsight-the ability to discriminate unseen stimuli in their clinically blind visual field defects when forced-choice procedures are used. Blindsight implies a sharp dissociation between visual performance and visual awareness, but signal detection theory indicates that it might be indistinguishable from the behavior of normal subjects near the lower limit of conscious vision, where the dissociations could arise trivially from using different response criteria during clinical and forced-choice tests. We tested the latter possibility with a hemianopic subject during yes-no and forced-choice detection of static and moving targets. His response criterion differed significantly between yes-no and forced-choice responding, and the difference was sufficient to produce a blindsight-like dissociation with bias-sensitive measures of performance. When measured independently of bias, his sensitivity to static targets was greater in the forced-choice than in the yes-no task (unlike normal control subjects), but his sensitivity to moving targets did not differ. Differences in response criterion could therefore account for dissociations between yes-no and forced-choice detection of motion, but not of static pattern. The results explain why patients with blindsight are apparently more often "aware" of moving stimuli than of static stimuli. However, they also imply that blindsight is unlike normal vision near threshold, and that pattern- and motion-detection in blindsight may depend on different sets of neural mechanisms during yes-no and forced-choice tests.     

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. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Magnetoencephalographic evidence for non-geniculostriate visual input to human cortical area V5

The aim of this study was to establish whether there is non-geniculostriate input to the extrastriate motion-sensitive area V5 in humans. Responses were measured with a SQUID neuro-magnetometer to motion stimuli presented within the blind hemifield of GY, a well-documented subject with a complete absence of the left primary visual cortical area V1. The motion stimulus was a 0.5c/deg, rapidly drifting (16Hz) achromatic sinusoidal grating. With this stimulus, the magnetic responses recorded over the temporo-parieto-occipital region in normals are well modelled by localized current sources in areas V1 and V5 (Anderson, S. J. et al., Proceedings of the Royal Society, London, Series B, 1996, 263, 423-431). As a control, evoked responses were measured to a 1.0 c/deg, stationary, photometrically isoluminant red/green sinusoidal grating. With the chromatic stimulus, the principal component of the magnetic responses recorded over the occipital pole in normals is well modelled by a current source in area V1 (Fylan, F. et al., Investigative Ophthalmology and Visual Science, 1995, 36, s1053). Both stimuli subtended 4 deg vertically by 6 deg horizontally, positioned such that the stimulus extended beyond the area of macular sparing into the lower field quadrant of the blind (or sighted) hemifield. Chromatic stimuli failed to evoked responses from GY's blind (contralateral) hemifield, consistent with there being no V1 activity in his left cortical hemisphere. However, motion stimuli did evoke responses from GY's blind hemifield, originating from a location consistent with activity in area V5. We further observed that both colour and motion stimuli evoked responses from GY's sighted (ipsilateral) hemifield. We conclude that there is non-geniculostriate input to extrastriate motion-sensitive areas in the human visual system, and that this pathway subserves the residual visual sensitivity to motion in the blind hemifield that has been demonstrated psychophysically in observer GY.     

A subcortical pathway to the right amygdala mediating "unseen" fear

Neuroimaging studies have shown differential amygdala responses to masked ("unseen") emotional stimuli. How visual signals related to such unseen stimuli access the amygdala is unknown. A possible pathway, involving the superior colliculus and pulvinar, is suggested by observations of patients with striate cortex lesions who show preserved abilities to localize and discriminate visual stimuli that are not consciously perceived ("blindsight"). We used measures of right amygdala neural activity acquired from volunteer subjects viewing masked fear-conditioned faces to determine whether a colliculo-pulvinar pathway was engaged during processing of these unseen target stimuli. Increased connectivity between right amygdala, pulvinar, and superior colliculus was evident when fear-conditioned faces were unseen rather than seen. Right amygdala connectivity with fusiform and orbitofrontal cortices decreased in the same condition. By contrast, the left amygdala, whose activity did not discriminate seen and unseen fear-conditioned targets, showed no masking-dependent changes in connectivity with superior colliculus or pulvinar. These results suggest that a subcortical pathway to the right amygdala, via midbrain and thalamus, provides a route for processing behaviorally relevant unseen visual events in parallel to a cortical route necessary for conscious identification.     

Topographical organization of inferior olive cells projecting to translation and rotation zones in the vestibulocerebellum of pigeons

Previous electrophysiological studies in pigeons have shown that the vestibulocerebellum can be divided into two parasagittal zones based on responses to optic flow stimuli. The medial zone responds best to optic flow resulting from self-translation, whereas the lateral zone responds best to optic flow resulting from self-rotation. This information arrives from the retina via a projection from the accessory optic system to the medial column of the inferior olive. In this study we investigated inferior olive projections to translational and rotational zones of the vestibulocerebellum using the retrograde tracer cholera toxin subunit B. Extracellular recordings of Purkinje cell activity (complex spikes) in response to large-field visual stimuli were used to identify the injection sites. We found a distinct segregation of inferior olive cells projecting to translational and rotational zones of the vestibulocerebellum. Translation zone injections resulted in retrogradely labeled cells in the ventrolateral area of the medial column, whereas rotation zone injections resulted in retrogradely labeled cells in the dorsomedial region of the medial column. Motion of any object through space, including self-motion of organisms, can be described with reference to translation and rotation in three-dimensional space. Our results show that, in pigeons, the brainstem visual systems responsible for detecting optic flow are segregated into channels responsible for the analysis of translational and rotational optic flow in the inferior olive, which is only two synapses from the retina.     

Disturbance of sleep in blindness

PURPOSE: To determine the prevalence and severity of sleep disturbance in blind subjects and its relation to the form and duration of visual loss. METHODS: Of 403 blind subjects (visual acuity of less than 20/200 or a visual field of less than 5 degrees) recruited for the study, 15 were excluded because of affective disorder as identified by Montgomery Asberg Depression Scale. The remaining 388 subjects and a comparison group of 44 normally sighted individuals underwent an interview, and the Pittsburgh Sleep Quality Index questionnaire was administered. Sleep disturbance was classified as mild, moderate, or severe. RESULTS: Disturbance of sleep was recorded in 189 (48.7%) of the blind subjects. The prevalence was higher and the sleep disturbance was more severe in those with no perception of light than in those with light perception or better visual acuity. In the comparison group, four (9.1%) had mild sleep disturbance only. The differences between blind subjects and normally sighted individuals were highly significant (P < .001). The most common sleep-related problem among the blind subjects was interrupted sleep, followed by increased sleep latency, short sleep duration, and daytime naps. Among the blind subjects, no correlation was found between the extent of sleep disturbance and the duration and pattern of visual loss. CONCLUSIONS: Blind subjects who retain light perception, as well as those with total loss of vision, have a high frequency of sleep disturbance, although disorder is more common and more severe in subjects with no light perception. Management of the sleep disturbance may improve the quality of life in the visually handicapped.     

Effects of inter- and intramodal selective attention to non-spatial visual stimuli: an event-related potential analysis

Event-related potentials (ERPs) were recorded to trains of rapidly presented auditory and visual stimuli. ERPs in conditions in which subjects attended to different features of visual stimuli were compared with ERPs to the same type of stimuli when subjects attended to different features of auditory stimuli. This design permitted us to study effects of variations in both intramodal and intermodal visual attention on the timing and topography of ERP components in the same experiment. There were no indications that exogenous N110, P140 and N180 components to line gratings of high and low spatial frequencies were modulated by either intra- or intermodal forms of attention. Furthermore, intramodal and intermodal attention effects on ERPs showed similar topographical distributions. These combined findings suggest that the same neural generators in extrastriate occipital areas are involved in both forms of attention. Visual ERPs elicited in the condition in which subjects were engaged in auditory selective attention showed a large positive displacement at the occipital scalp sites relative to ERPs to attended and unattended stimuli in the visual condition. The early onset of this positivity might be associated with a highly confident and early rejection of the irrelevant visual stimuli, when these stimuli are presented among auditory stimuli. In addition, the later onset of selection potentials in the intramodal condition suggests that a more precise stimulus selection is needed when features of visual stimuli are rejected among other features of the same stimulus pattern, than when visual stimuli are rejected among stimuli of another modality.     

The directional effects of passive eye movement on the directional visual responses of single units in the pigeon optic tectum

We have investigated the visual responses of 184 single units located in the superficial layers of the optic tectum (OT) of the decerebrate, paralysed pigeon. Visual responses were similar to those reported in non-decerebrate preparations; most units responded best to moving visual stimuli, 18% were directionally selective (they had a clear preference for a particular direction of visual stimulus movement), 76% were plane-selective (they responded to movement in either direction in a particular plane). However, we also found that a high proportion of units showed some sensitivity to the orientation of visual stimuli. We examined the effects of extraocular muscle (EOM) afferent signals, induced by passive eye movement (PEM), on the directional visual responses of units. Visual responses were most modified by particular directions of eye movement, although there was no unique relationship between the direction of visual stimulus movement to which an individual unit responded best and the direction of eye movement that caused the greatest modification of that visual response. The results show that EOM afferent signals, carrying information concerning the direction of eye movement, reach the superficial layers of the OT in the pigeon and there modify the visual responses of units in a manner that suggests some role for these signals in the processing of visual information.     

Episodic multiregional cortical coherence at multiple frequencies during visual task performance

The way in which the brain integrates fragmentary neural events at multiple locations to produce unified perceptual experience and behaviour is called the binding problem. Binding has been proposed to involve correlated activity at different cortical sites during perceptuomotor behaviour, particularly by synchronization of narrow-band oscillations in the gamma-frequency range (30-80 Hz). In the rabbit olfactory system, inhalation induces increased gamma-correlation between sites in olfactory bulb and cortex. In the cat visual system, coherent visual stimuli increase gamma-correlation between sites in both the same and different visual cortical areas. In monkeys, some groups have found that gamma-oscillations transiently synchronize within striate cortex, superior temporal sulcus and somatosensorimotor cortex. Others have reported that visual stimuli produce increased broad-band power, but not gamma-oscillations, in several visual cortical areas. But the absence of narrow-band oscillations in itself does not disprove interregional synchronization, which may be a broad-band phenomenon. We now describe episodes of increased broad-band coherence among local field potentials from sensory, motor and higher-order cortical sites of macaque monkeys performing a visual discrimination task. Widely distributed sites become coherent without involving other intervening sites. Spatially selective multiregional cortical binding, in the form of broad-band synchronization, may thus play a role in primate perceptuomotor behaviour.     

The Von Restorff effect in visual object recognition memory in humans and monkeys. The role of frontal/perirhinal interaction

This study reports the development of a new, modified delayed matching to sample (DMS) visual recognition memory task that controls the relative novelty of test stimuli and can be used in human and nonhuman primates. We report findings from normal humans and unoperated monkeys, as well as three groups of operated monkeys. In the study phase of this modified paradigm, subjects studied lists of two-dimensional visual object stimuli. In the test phase each studied object was presented again, now paired with a new stimulus (a foil), and the subject had to choose the studied item. In some lists one study item (the novel or isolate item) and its associated foil differed from the others (the homogenous items) along one stimulus dimension (color). The critical experimental measure was the comparison of the visual object recognition error rates for isolate and homogenous test items. This task was initially administered to human subjects and unoperated monkeys. Error rates for both groups were reliably lower for isolate than for homogenous stimuli in the same list position (the von Restorff effect). The task was then administered to three groups of monkeys who had selective brain lesions. Monkeys with bilateral lesions of the amygdata and fornix, two structures that have been proposed to play a role in novelty and memory encoding, were similar to normal monkeys in their performance on this task. Two further groups--with disconnection lesions of the perirhinal cortex and either the prefrontal cortex or the magnocellular mediodorsal thalamus--showed no evidence of a von Restorff effect. These findings are not consistent with previous proposals that the hippocampus and amygdala constitute a general novelty processing network. Instead, the results support an interaction between the perirhinal and frontal cortices in the processing of certain kinds of novel information that support visual object recognition memory.     

Rats spontaneously discriminate purely visual, two-dimensional stimuli in tests of recognition memory and perceptual oddity.

Animal models have been central to advances made in understanding the neural basis of human cognition, but maximizing the use of animal models requires tasks that match those used to assess human subjects. Tasks used in humans frequently use visual 2-dimensional stimuli, assess 1-trial learning, and require little pretraining. This article describes novel versions of 2 tasks for the rat, spontaneous object recognition and spontaneous oddity preference, both of which use purely visual, 2-dimensional picture-card stimuli, test 1-trial learning, and require no pretraining. Rats showed robust memory for a variety of picture-card stimuli, demonstrating almost no loss of memory for some of the stimulus types even after a 2-hr delay period. Rats were able to show spontaneous oddity preference for all 3 visual stimulus types tested (photos, shapes, and patterns), as well as for 3-dimensional objects. These 2 tasks are quick to administer, involve no fearful learning associations, and require a simple apparatus. They may be particularly useful for high-throughput pharmacological or genetic screening using rodent models. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Imagery limitations in totally congenitally blind subjects.

Research on totally blind subjects performing tasks that involve visual imagery has often shown that they do not behave differently from matched sighted subjects, even when their blindness is congenital. If visual imagery is based on visual perception, such tasks may not require visual imagery. In the present article visual images are considered as representations maintaining some properties of visible objects and constructed on the basis of information from various sources. Owing to the absence of visual experience, the limitations of such representations are explored in a series of experiments requiring memorization of single nouns, pairs of nouns, or triplets of nouns associated with a cue noun. Recall by blind subjects was impaired when multiple interactive images (with noun pairs and triplets) are formed. The poorer recall of blind subjects reflected also loss of order information. Recall was better for both groups with locative noun cues and high-imagery targets. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Retinotopic organisation of cortical mechanisms responsive to colour: evidence from patient studies

This paper deals with the visual responses of three patients who have impaired colour vision consequent on cortical dysfunction which, in two of them, is associated with demonstrable neuronal damage. The studies to be described are concerned particularly with the spatial attributes of their chromatic response mechanisms. Data are presented which establish that a hemianope GY has coarse chromatic discrimination for large stimuli located within his 'blind' hemifield. GY responds to stimuli containing differently coloured equiluminant components as if the coloured components were averaged over the whole field and it is speculated that such spatial averaging may correspond to the process which, in normal vision, provides compensation for change of illuminant in order to achieve colour constancy. Colour constancy is impaired in a second patient, BL, who has cortical lesions involving the lingual and fusiform gyri, areas which are partially spared in GY. It is shown that movement, but not colour, presented to GY's normal hemifield generates a response localised in his blind hemifield and disinhibitory interaction between movement and colour is illustrated for a patient MW, in whom colour chromatic stimuli generate spreading inhibition of visual responses. This inhibitory interaction is propagated between widely separated stimuli, including those which are located on opposite sides of the vertical meridian. We discuss these experimental results in relation to anatomical and physiological mechanisms of the primate visual cortex.     

Stimulus features eliciting visual responses from neurons in the nucleus lentiformis mesencephali in pigeons

The purpose of the present study was to find out what particular stimulus features, in addition to the direction and velocity of motion, specifically activate neurons in the nucleus lentiformis mesencephali (nLM) in pigeons. Visual responses of 60 nLM cells to a variety of computer-generated stimuli were extracellularly recorded and quantitatively analyzed. Ten recording sites were histologically verified to be localized within nLM with cobalt sulfide markings. It was shown that the pigeon nLM cells were specifically sensitive to the leading edge moving at the optimal velocity in the preferred direction through their excitatory receptive fields (ERFs). Generally speaking, nLM cells preferred black edges to white ones. However, this preference cannot be explained by OFF-responses to a light spot. The edge sharpness was also an essential factor influencing the responsive strength, with blurred edges producing little or no visual responses at all. These neurons vigorously responded to black edge orientated perpendicular to, and moved in, the preferred direction; the magnitude of visual responses was reduced with changing orientation. The spatial summation occurred in all neurons tested, characterized by the finding that neuronal firings increased as the leading edge was lengthened until saturation was reached. On the other hand, it appeared that nLM neurons could not detect any differences in the shape and area of stimuli with an identical edge. These data suggested that feature extraction characteristics of nLM neurons may be specialized for detecting optokinetic stimuli, but not for realizing pattern recognition. This seems to be at least one of the reasons why large-field gratings or random-dot patterns have been used to study visual responses of accessory optic neurons and optokinetic nystagmus, because many high-contrast edges in these stimuli can activate a neuron to periodically discharge, or groups of neurons to simultaneously fire to elicit optokinetic reflex.     

Nystagmus in periventricular leucomalacia

BACKGROUND/AIMS: Periventricular leucomalacia (PVL) is a lesion in the immature brain involving the optic radiation. Children with PVL have visual problems including crowding, visual field defects, strabismus, and visual perceptual/cognitive deficits, together with nystagmus. They often have optic nerve hypoplasia seen either as small discs or as large cupping of normal sized optic discs. This study aimed to perform eye movement recordings in a group of children with PVL in order to characterise and classify the nystagmus. METHODS: 19 children with PVL on cerebral imaging underwent eye movement recordings with the Ober-2 infrared reflection technique. RESULTS: 16 of the 19 subjects had horizontal nystagmus. CONCLUSION: The present study shows that nystagmus is commonly seen in children with PVL.     

Single units and conscious vision

Figures that can be seen in more than one way are invaluable tools for the study of the neural basis of visual awareness, because such stimuli permit the dissociation of the neural responses that underlie what we perceive at any given time from those forming the sensory representation of a visual pattern. To study the former type of responses, monkeys were subjected to binocular rivalry, and the response of neurons in a number of different visual areas was studied while the animals reported their alternating percepts by pulling levers. Perception-related modulations of neural activity were found to occur to different extents in different cortical visual areas. The cells that were affected by suppression were almost exclusively binocular, and their proportion was found to increase in the higher processing stages of the visual system. The strongest correlations between neural activity and perception were observed in the visual areas of the temporal lobe. A strikingly large number of neurons in the early visual areas remained active during the perceptual suppression of the stimulus, a finding suggesting that conscious visual perception might be mediated by only a subset of the cells exhibiting stimulus selective responses. These physiological findings, together with a number of recent psychophysical studies, offer a new explanation of the phenomenon of binocular rivalry. Indeed, rivalry has long been considered to be closely linked with binocular fusion and stereopsis, and the sequences of dominance and suppression have been viewed as the result of competition between the two monocular channels. The physiological data presented here are incompatible with this interpretation. Rather than reflecting interocular competition, the rivalry is most probably between the two different central neural representations generated by the dichoptically presented stimuli. The mechanisms of rivalry are probably the same as, or very similar to, those underlying multistable perception in general, and further physiological studies might reveal much about the neural mechanisms of our perceptual organization.     

Cortical spreading depression induces an LTP-like effect in rat neocortex in vitro

We studied the auditory neurons in the optic tectum of the unanesthetized pigeon, using single-unit recordings and acoustic free-field stimulation. Most units showed spatial tuning, with best areas located in the contralateral hemifield. All units responded also to visual stimuli, the auditory best areas being in rough alignment with visual receptive fields.     

Laser scanning tomography of the optic nerve head in ocular hypertension and glaucoma

BACKGROUND: This study evaluated the ability of laser scanning tomography to distinguish between normal and glaucomatous optic nerve heads, and between glaucomatous subjects with and without field loss. METHODS: 57 subjects were classified into three diagnostic groups: subjects with elevated intraocular pressure, normal optic nerve heads, and normal visual fields (n = 10); subjects with glaucomatous optic neuropathy and normal visual fields (n = 30); and subjects with glaucomatous optic neuropathy and repeatable visual field abnormality (n = 17). Three 10 degrees image series were acquired on each subject using the Heidelberg retina tomograph (HRT). From the 14 HRT stereometric variables, three were selected a priori for evaluation: (1) volume above reference (neuroretinal rim volume), (2) third moment in contour (cup shape), and (3) height variation contour (variation in relative nerve fibre layer height at the disc margin). Data were analysed using analysis of covariance, with age as the covariate. RESULTS: Volume above reference, third moment in contour, and mean height contour were significantly different between each of the three diagnostic groups (p < 0.001). Height variation contour showed no significant difference among the three diagnostic groups (p = 0.906). CONCLUSIONS: The HRT variables measuring rim volume, cup shape, and mean nerve fibre layer height distinguished between (1) subjects with elevated intraocular pressures and normal nerve heads, and glaucomatous optic nerve heads, and (2) glaucomatous optic nerve heads with and without repeatable visual field abnormality. This study did not directly assess the ability of the HRT to identify patients at risk of developing glaucoma. It is hypothesised that the greatest potential benefit of laser scanning tomography will be in the documentation of change within an individual over time.     

When seeing doesn't matter: Assessing the after-effects of tactile distractor processing in the blind and the sighted.

Negative priming (NP) refers to the finding that people's responses to probe targets previously presented as prime distractors are usually slower than to unrepeated stimuli. Intriguingly, the effect sizes of tactile NP were much larger than the effect sizes for visual NP. We analyzed whether the large tactile NP effect is just a side effect of the higher difficulty when processing tactile compared to visual stimuli. Thus, we analyzed tactile NP in a sample of blind participants and in a control sample of sighted participants. Although the blind participants handled the tactile stimuli with ease, we found no evidence that the size of the tactile NP effect diminished. In two control experiments with sighted participants, we varied the processing difficulty in the visual and tactile modality and found that both modality and processing difficulty had an effect on the size of NP. Taken together, our data show that the difficulty associated with processing tactile stimuli is only partially the reason for the unusual large tactile NP effect. These results suggest that non-spatial tactile distractors are processed and selected quite differently from visual distractors. (PsycINFO Database Record (c) 2011 APA, all rights reserved)