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Biomedical subjects

J L Barbur

Publications and source records attributed to J L Barbur.

At least 19 recordsLinked to original sources

The unseen color aftereffect of an unseen stimulus: insight from blindsight into mechanisms of color afterimages.

We show here that, in the absence of a direct geniculostriate input in human subjects, causing loss of sight in the visual half-field contralateral to the damage, the pupil responds selectively to chromatic modulation toward the long-wavelength (red) region of the spectrum locus even when the stimulus is isoluminant for both rods and cones and entirely restricted to the subjects' "blind" hemifields. We also show that other colors are less or wholly ineffective. Nevertheless, red afterimages, generated by chromatic modulation toward the green region of the spectrum locus, also cause constrictions of the pupil even when green stimuli are themselves completely ineffective in the blind hemifield. Moreover, human subjects with damage to or loss of V1 are typically completely unaware of the stimulus that generates the aftereffect or of the aftereffect itself, both of which can be seen clearly in normal vision. The results show that pupillary responses can reveal the processing of color afterimages in the absence of primary visual cortex and in the absence of acknowledged awareness. This phenomenon is therefore a striking example of "blindsight" and makes possible the formulation of a model that predicts well the observed properties of color afterimages.

Adult

Differential pupillary constriction and awareness in the absence of striate cortex.

The fact that the pupil constricts differentially to visual stimuli in the absence of changes in light energy makes it a valuable tool for studying normal function as well as residual capacity in hemianopic subjects. When pupillometrically effective stimuli such as equiluminant gratings or coloured patches with an abrupt onset and offset are presented to the 'blind' hemifield, a hemianopic subject with damage largely restricted to striate cortex (V1) sometimes reports being 'aware' of the transient onset/offset, although without 'seeing' as such. The question addressed here is whether the pupil still responds in the condition of blindsight in its strict sense--i.e. discriminative capacity in the absence of acknowledged awareness--when stimuli are deliberately designed to eliminate awareness. This was accomplished by making stimulus onset and offset slow and gradual. The results with a well-studied hemianope, G.Y., demonstrate that there is still a pupillary constriction to isoluminant achromatic gratings and red-coloured stimuli, although reduced in size, in the absence of acknowledged awareness.

Accidents, Traffic

Visual processing levels revealed by response latencies to changes in different visual attributes.

Visual latencies, and their variation with stimulus attributes, can provide information about the level in the visual system at which different attributes of the image are analysed, and decisions about them made. A change in the colour, structure or movement of a visual stimulus brings about a highly reproducible transient constriction of the pupil that probably depends on visual cortical mechanisms. We measured this transient response to changes in several attributes of visual stimuli, and also measured manual reaction times to the same stimulus changes. Through analysis of latencies, we hoped to establish whether changes in different stimulus attributes were processed by mechanisms at the same or different levels in the visual pathway. Pupil responses to a change in spatial structure or colour are almost identical, but both are ca. 40 ms slower than those to a change in light flux, which are thought to depend largely on subcortical pathways. Manual reaction times to a change in spatial structure or colour, or to the onset of coherent movement, differ reliably, and all are longer than the reaction time to a change in light flux. On average, observers take 184 ms to detect a change in light flux, 6 ms more to detect the onset of a grating, 30 ms more to detect a change in colour, and 37 ms more to detect the onset of coherent motion. The pattern of latency variation for pupil responses and reaction times suggests that the mechanisms that trigger the responses lie at different levels in cortex. Given our present knowledge of visual cortical organization, the long reaction time to the change in motion is surprising. The range of reaction times across different stimuli is consistent with decisions about the onset of a grating being made in V1 and decisions about the change in colour or change in motion being made in V4.

Adult

Pupil responses to stimulus color, structure and light flux increments in the rhesus monkey.

Visual stimuli that isolate pupil color and pupil grating responses in human vision have been used to investigate the properties, of stimulus-specific pupil responses in the rhesus monkey. We measured and compared pupil responses to light flux increments, isoluminant chromatic stimuli, and gratings of equal and lower space-averaged luminance. The parameters investigated were luminance contrast and chromatic saturation. The results demonstrate clearly the existence of pupil color, pupil grating and pupil light reflex responses in the rhesus monkey. Comparison of pupil color and pupil grating responses of equivalent amplitude reveals similar onset response latencies. However, both are approximately 40 ms longer than the corresponding pupil light reflex latency. In general these pupil responses are qualitatively similar to those observed in humans. However, when compared to equivalent human data, pupil onset response latencies are some 80-100 ms shorter and the pupil shows more rapid recovery from constriction.

Adult

Awareness and confidence ratings in motion perception without geniculo-striate projection.

We have previously reported that under certain experimental conditions, a subject with damaged primary visual cortex performed significantly above chance in discriminating motion direction and orientation either with or without awareness of the stimulus presentation in his blind hemifield. The subject's performance varied as a function of stimulus speed, excursion length, and background luminance. Present observations confirm the previous findings of above chance discrimination scores both with and without awareness, but nevertheless indicate an overall increase in sensitivity over the past 2 years. In addition to discriminating the direction of motion, the subject was asked on every trial to report either awareness or confidence or both, on a six-point scale. The results show that the introduction of a six-point scale for the reported level of awareness yielded similar results to those when the subject was given a binary choice to indicate the presence or absence of conscious awareness. The area under the receiver operating characteristic (ROC) curves obtained from the rating data was used to compare results across different experimental conditions. It was found that although the subject's reported confidence and awareness varied monotonically as a function of stimulus speed, they were not equivalent measures.

Adult

The effect of age on the light scattering characteristics of the eye.

The rising mean age of our population has increased the need for understanding the physiologic consequences of ageing on visual function. Changes due to age were evaluated using a new scatter test implemented on the P_SCAN 100 pupillometer apparatus (Barbur, 1991). The test yields the full scatter function of the eye and also permits simultaneous measurement of pupil size (Barbur et al., 1995). In addition, contrast sensitivity was measured using sine wave gratings. The 28 subjects had a spherical refractive error between +0.50 DS and -0.25 DS, and astigmatism of less than -0.50 DC, V.A. of at least 6/6, and were ophthalmologically normal. To facilitate statistical analysis, subjects were classified into five groups according to age. For younger subjects (under 45 years), k', the integrated straylight parameter, varied little with age. For this group, k' ranged from 4.9 to 8.1. For subjects aged over 45 years, k' increased with age, ranging from 10.7 to 19.7. One way analysis of variance showed Group 5 (60 year olds) to have significantly greater k' than 20, 30, 40 and 50 year olds (P = 0.000). A slight downward shift in the contrast sensitivity function was seen over the age of 45. Significant differences between older and younger subjects were found at spatial frequencies of 3, and 10 cpd (P = 0.081, P = 0.002, respectively). Pupil diameter was found to reduce with age, but there was no significant difference between groups. Therefore, above 45 years, the ageing eye reveals a more rapid increase in forward scatter, and a reduction in contrast sensitivity, despite apparently good visual acuity.

Adolescent

Residual processing of chromatic signals in the absence of a geniculostriate projection.

We have investigated the residual processing of chromatic signals in a subject with unilateral damage to the primary visual cortex using psychophysical, pupillometric and functional magnetic resonance imaging (fMRI) methods. Of particular interest was to establish the correlation between the subject's ability to make use of chromatic signals in the blind hemifield to discriminate between different coloured targets, the corresponding residual pupil colour responses and the level and location of cortical activation generated by the same stimuli as revealed by fMRI. The results obtained using the three different experimental approaches are consistent and suggest that retrograde degeneration of thalamic and retinal chromatic processing mechanisms caused by damage to the primary visual cortex in man does not abolish completely the ability to process chromatic signals particularly when large, long-wavelength stimuli are employed.

Color Perception

Pattern of neuronal activity associated with conscious and unconscious processing of visual signals.

Following striate cortex damage in monkeys and humans there can be residual function mediated by parallel visual pathways. In humans this can sometimes be associated with a "feeling" that something has happened, especially with rapid movement or abrupt onset. For less transient events, discriminative performance may still be well above chance even when the subject reports no conscious awareness of the stimulus. In a previous study we examined parameters that yield good residual visual performance in the "blind" hemifield of a subject with unilateral damage to the primary visual cortex. With appropriate parameters we demonstrated good discriminative performance, both with and without conscious awareness of a visual event. These observations raise the possibility of imaging the brain activity generated in the "aware" and the "unaware" modes, with matched levels of discrimination performance, and hence of revealing patterns of brain activation associated with visual awareness. The intact hemifield also allows a comparison with normal vision. Here we report the results of a functional magnetic resonance imaging study on the same subject carried out under aware and unaware stimulus conditions. The results point to a shift in the pattern of activity from neocortex in the aware mode, to subcortical structures in the unaware mode. In the aware mode prestriate and dorsolateral prefrontal cortices (area 46) are active. In the unaware mode the superior colliculus is active, together with medial and orbital prefrontal cortical sites.

Brain

Pupil response triggered by the onset of coherent motion.

BACKGROUND: Recent studies have shown that transient constrictions of the pupil can be elicited by visual stimuli that do not cause an increment in light flux level on the retina. Such stimuli include achromatic gratings and isoluminant chromatic patterns. METHOD: We investigated pupillary responses to the onset of coherent movement generated in a pattern of dots in random motion. Measurements were carried out in normal observers and in a subject with hemianopia caused by damaged primary visual cortex. RESULTS: The experimental findings show that the onset of coherent motion triggers systematic constrictions of the pupil that cannot be accounted for in terms of a pupil light reflex response. We labelled these constrictions Pupil motion responses (PMRs). Results show that PMRs have large response latencies and on average are of small response amplitudes. The dependence of PMRs on changes in motion parameters such as stimulus speed and direction of motion has been investigated. CONCLUSIONS: The existence of PMRs to the onset of the coherent motion in human vision has been demonstrated. These new findings are discussed in relation to the psychophysical and physiological data on motion perception and the possible pathways involved in the control of the pupil response.

Hemianopsia

Motion discrimination of single targets: comparison of preliminary findings in normal subjects and patients with glaucoma.

BACKGROUND: Luminance, global motion and flicker sensitivities are affected in patients with primary open-angle glaucoma. Although no theoretical model has been put forward to explain the observed reduction in sensitivity in this patient group, these findings have often been attributed to diffuse and selective loss of large optic nerve fibres. METHODS: Movement processing was investigated using an optical projection system that generates smooth, continuous motion with control of speed, displacement and motion direction. Motion-displacement and direction-discrimination thresholds were measured in eight normal subjects and in three patients with diagnosed glaucoma. At each speed tested, targets were presented for a range of displacements and thresholds were extracted after probit analysis. The measurements were carried out both foveally and at 19 degrees in the periphery and provided the data necessary to develop and optimise a model of motion perception based on multiple time delays for the correlation of signals that map progressively more distant parts of the visual field. RESULTS: Our preliminary findings show that direction discrimination can be at chance level even for large displacements when motion is detected 80% of the time. Model simulations show that specific changes in the spatial sampling interval and the speed of transmission of the motion signals involved can cause the observed reduction in motion sensitivity and direction discrimination in patients with glaucoma. CONCLUSIONS: A model for motion detection and direction discrimination of single targets has been proposed to account for the measured functional relationship between motion displacement thresholds and target speed in normal subjects. Tested patients with glaucoma show reduced motion sensitivity and poor discrimination of motion direction. The type of degraded performance observed experimentally in glaucoma patients is also predicted by the model. Such predictions require specific changes in model parameters that may be indicative of changes in the retina caused by the disease.

Adult

Parameters affecting conscious versus unconscious visual discrimination with damage to the visual cortex (V1).

When the visual (striate) cortex (V1) is damaged in human subjects, cortical blindness results in the contralateral visual half field. Nevertheless, under some experimental conditions, subjects demonstrate a capacity to make visual discriminations in the blind hemifield (blindsight), even though they have no phenomenal experience of seeing. This capacity must, therefore, be mediated by parallel projections to other brain areas. It is also the case that some subjects have conscious residual vision in response to fast moving stimuli or sudden changes in light flux level presented to the blind hemifield, characterized by a contentless kind of awareness, a feeling of something happening, albeit not normal seeing. The relationship between these two modes of discrimination has never been studied systematically. We examine, in the same experiment, both the unconscious discrimination and the conscious visual awareness of moving stimuli in a subject with unilateral damage to V1. The results demonstrate an excellent capacity to discriminate motion direction and orientation in the absence of acknowledged perceptual awareness. Discrimination of the stimulus parameters for acknowledged awareness apparently follows a different functional relationship with respect to stimulus speed, displacement, and stimulus contrast. As performance in the two modes can be quantitatively matched, the findings suggest that it should be possible to image brain activity and to identify the active areas involved in the same subject performing the same discrimination task, both with and without conscious awareness, and hence to determine whether any structures contribute uniquely to conscious perception.

Adult

Insights into the different exploits of colour in the visual cortex.

A new method that allows controlled masking of luminance contrast has been developed to study the use of chromatic signals in human vision. The method also makes it possible to examine the different uses of chromatic signals (e.g. the generation of perceived colour, or the construction and representation of object structure and form). By using this technique, we studied the threshold detection of chromatic signals in normal trichromats. The results show that chromatic signals are virtually unaffected by ongoing, randomly varying, luminance contrast changes. These findings suggest that chromatic signals are either processed independently or can be separated completely from any confounding luminance contrast components in the stimulus. Thresholds for detection of colour changes only, and for extraction of stimulus structure from chromatic signals in normal trichromats, in subjects with single cone receptor deficiency (i.e. dichromats) and in three subjects with abnormal colour vision caused by bilateral damage to ventromedial, extra-striate visual cortex (i.e. subjects with cerebral achromatopsia) have also been measured. No significant difference in thresholds for the two conditions was observed either in normal trichromats or in dichromats. Subjects with cerebral achromatopsia, however, reveal markedly different thresholds. The results suggest that chromatic signals are processed independently to generate perceived object colour or to construct spatially structured objects, and that these functions involve different neural substrates. The results help to explain, at least in part, why cerebral achromatopsia is a heterogeneous disorder, and why there can be significant differences in the effective use of chromatic signals in subjects described as cerebral achromatopsics.

Color Perception

Spatial and temporal response properties of residual vision in a case of hemianopia.

Residual vision in subjects with damage of the primary visual cortex (striate cortex) has been demonstrated in many previous studies and is taken to reflect the properties of known subcortical and extrastriate visual pathways. In this report we describe psychophysical experiments carried out on a subject clinically blind in half of his visual field (i.e. homonymous hemianopia) caused by striate cortex damage. They reveal the existence of two distinct channels mediating such vision. One channel responds to spatial structure and the other to light flux changes. The spatially tuned channel has a peak response at about 1.2 cycles per degree and shows rapid loss of sensitivity at both high and low spatial frequencies. This channel does not respond to diffuse illumination. The light flux channel, however, responds only to sudden increments in light flux levels on the retina and shows extensive spatial summation. Both channels require transient inputs, with a peak sensitivity at about 10 cycles per second and show virtually complete attenuation at temporal frequencies below 2 cycles per second. The spatiotemporal characteristics of these two channels account for much of the reported limits of visual performance attributed to subcortical or extrastriate pathways in some patients, and especially for their relatively good sensitivity for the detection of abrupt, transient stimuli or fast-moving targets. A new method is also applied to the measurement of the amount of light scatter in the eye. The measurements show that light scatter into the sighted hemifield could not account for the results obtained with the stimuli used to characterized the residual vision of this subject.

Accidents, Traffic

Pupillary function in human amblyopia.

Quantitative measurements of pupillary function (response amplitude and latency) were made for normal eyes and for normal and fellow amblyopic eyes of groups of strabismic and anisometropic amblyopes. Stimuli consisted of luminance modulation of a large, evenly lit area (pupil light reflex) as well as contrast modulation of sinusoidal gratings (pupil grating response) of fixed, space-averaged luminance. Measurements were made of the direct and the consensual reflex under monocular stimulation. A comparison of the amplitude of the pupil light reflex as a function of luminance modulation showed no significant differences between normal and fellow amblyopic eyes for both the strabismic and anisometropic groups of amblyopes studied. A similar comparison of the associated response latencies showed significant difference between normal and fellow amblyopic eyes for both groups. In general, reductions in response amplitude and latency of the pupil grating response were found in individuals from each group when comparing the good and the affected eyes, although the observed group differences were only significant in the strabismic group. Interestingly, statistically significant reductions in both amplitude and latency for both the pupil light reflex and the pupil grating response were found between the eyes of normal observers and the so-called normal eyes of amblyopes in both groups studied. These results suggest that the type of pupillary deficit in amblyopia is a complicated one, depending not only on the type of amblyopia (strabismic or anisometropic) and the type of stimulus employed (light or pattern), but also on the parameter assessed (amplitude or latency) and whether the amblyopic result is referenced to its fellow normal eye or to the normal eye of a non-amblyopic observer. Since the pupil response to light flux changes is not mediated exclusively via the retinal projection to the midbrain and may also involve the activity of central visual pathways, the results obtained in this study cannot be used to provide definitive evidence for the site of abnormality in amblyopia.

Adolescent

Conscious visual perception without V1.

We used the technique of PET to determine whether visual signals reach visual area V5, specialized for visual motion, when a human patient, blinded by a lesion in area V1, discriminates the direction of motion of visual stimuli and shows, through his verbal reports, that he is consciously aware of both the nature of the visual stimulus and its direction of motion. The results showed that area V5 was active without a parallel activation of area V1, implying that the visual input can reach V5 without passing first through V1 and that such an input is sufficient for both the discrimination and the conscious awareness of the visual stimulus.

Adult

New method based on random luminance masking for measuring isochromatic zones using high resolution colour displays.

A new method of measuring normal hue discrimination ellipses and dichromatic zones using a high resolution colour monitor is described. The test involves the detection of chromatic bars on a grey background (x = 0.305, y = 0.323) having a luminance of 34 cd m-2. Elements of the background matrix of square checks are varied randomly in luminance in space and time to provide random luminance masking (RLM) which compensates for differences in the relative luminous efficiency of different observers. The measurement technique provides a rapid and comprehensive colour vision test. Typical results are presented for normal trichromats, protanopes and deuteranopes without RLM and with the RLM set of 25%. The size of the discrimination ellipse in normal observers is the same in both viewing conditions, but the use of the RLM technique reveals the extent of the isochromatic zones in colour deficient observers.

Color Perception Tests

Pupillary responses to stimulus structure, colour and movement.

Pupillary responses to stimuli which favour the preferential stimulation of neural mechanisms involved in the detection of visual attributes such as colour, spatial structure, movement and light flux changes on the retina have been measured and compared. Pupil responses to a decrement in stimulus luminance (i.e., a flash of darkness), suggest that at least three components are involved in this response, their relative contribution being determined largely by stimulus size, contrast and presentation time. A comparison of pupil responses to gratings of equal and lower space-averaged luminance shows that the amplitude of pupillary constriction at grating onset for the equal luminance condition is about twice that measured with similar gratings in the lower luminance condition. Pupillary responses to chromatic isoluminant gratings are in general of longer latency when compared to responses of similar amplitude elicited by achromatic gratings. Small pupillary constrictions elicited by the onset of coherent movement in dynamic, random dot patterns are also demonstrated under stimulus conditions which eliminate pupillary responses to sudden light flux changes on the retina. The results support an earlier hypothesis which suggests that the onset of sudden changes in neural activity in the visual cortex when a visual stimulus is presented to the eye causes an overall perturbation which weakens transiently the regulatory inhibitory input to the pupillomotor nucleus. This, in turn, results in a transient increase in the efferent parasympathetic innervation of the iris sphincter muscle and hence the observed constriction of the pupil. The characteristics of the pupillary response reflect the properties of the mechanisms and the number of neurones which participate in the detection of each stimulus attribute.

Color Perception

Factors affecting visual sensitivity in a hemianopic subject.

A well-studied subject with visual cortex damage (G.Y.) was tested in his hemianopic field with temporally modulated sinusoidal and square-wave gratings. The purpose was to use an extended range of parameters to obtain a detailed spatiotemporal specification of his residual vision and to try to resolve the discrepancy between negative findings of Hess and Pointer (1989) and previous positive claims. Both the spatial and temporal parameters could be Gaussian-weighted. Detection as a function of spatial frequency, contrast, temporal modulation frequency, stimulus size, and slope of the temporal and spatial Gaussian functions was investigated using a two-alternative forced-choice procedure. The most important parameters for this subject were found to be the slope of the temporal Gaussian function and the size and contrast of gratings. With optimum parameters he could reliably achieve a score of 95-100% correct in his 'blind' field. The results are consistent with earlier studies of this subject, especially his ability to respond to moving stimuli, and also may account for why negative results had been reported for him when particular fixed parameters were used.

Contrast Sensitivity