Search PubMed⌕ Search

Biomedical subjects

Alan Cowey

Publications and source records attributed to Alan Cowey.

At least 19 recordsLinked to original sources

Auditory evoked visual awareness following sudden ocular blindness: an EEG and TMS investigation.

Neurophysiological and neuroanatomical studies have provoked controversy about whether the visual cortex may be more modifiable than previously believed. Auditory processing is enhanced in blind compared to sighted people, and the enhancement might reflect encroachment of auditory transmission onto visual cortex. To address this issue, we recorded the auditory event-related potentials (ERPs) correlated with auditory related paradoxical visual awareness in a subject with traumatic total late-onset blindness. We found that (1) there was auditory related brain activity over the occipital visual scalp regions starting from a very early stage (< 80 ms) and (2) this occipital activity was significantly different between "visually aware" and "visually unaware" responses in the P1 (40-80 ms) component following meaningful stimuli. There was also a significant difference between responses with and without visual awareness in the N1 (100-120 ms) component following either tones or meaningful stimuli. The phosphenes accompanying auditory stimuli in the ERP experiment were always perceived to be directly in front of the subject and this was reproduced by transcranial magnetic stimulation over the blind primary visual cortex and by sudden sounds delivered to the side or behind the subject. The TMS induced phosphenes were restricted to the central part of the space and were, at least qualitatively, the same as those induced by sounds. The results are clear evidence that human perceptual functions can be reorganized after sudden, late-onset, total ocular blindness.

Acoustic Stimulation↗

TMS can reveal contrasting functions of the dorsal and ventral visual processing streams.

In order to investigate the functional specificity of the dorsal and ventral visual processing steams we used transcranial magnetic stimulation (TMS) to briefly disrupt one or the other while subjects performed three tasks, involving discrimination of colour or shape or relative position. TMS was delivered over right posterior parietal cortex (PPC) or right lateral occipital (LO) cortex, regions known to have visuo-spatial and object processing properties respectively. LO but not PPC stimulation had a significant effect on reaction time when subjects were asked to make a discrimination of relative shape. PPC stimulation had a significant effect when subjects were asked to discriminate relative position of the same shapes. Stimulation of LO also lengthened reaction times on the position task. There were no effects of stimulation at either site on colour discrimination. Results are discussed within the framework of how the dorsal stream and ventral stream are dissociated following their damage in neurological patients and possible ways in which they may interact in the normal brain.

Adult↗

The role of human extra-striate visual areas V5/MT and V2/V3 in the perception of the direction of global motion: a transcranial magnetic stimulation study.

Several published single case studies reveal a double dissociation between the effects of brain damage in separate extra-striate cortical visual areas on the perception of global visual motion defined by a difference in luminance (first-order motion) versus motion defined by a difference in contrast (second-order motion). In particular, the medial extrastriate cortical region V2/V3 seems to be crucial for the perception of first-order motion, but not for second-order, whereas a lateral and more anterior portion of the cortex close to the temporo-parieto-occipital junction (in the territory of the human motion area hV5/MT+) seems to be essential only for the perception of second-order motion. In order to test the hypothesis of a functional specialization of different visual areas for different types of motion, we applied repetitive transcranial magnetic stimulation (rTMS) unilaterally over areas V2/V3, V5/MT, or posterior parietal cortex (PPC) while subjects performed a 2AFC task with first- or second-order global motion displays in the contralateral visual field. Results showed a comparable disruption of the two types of motion, with both rTMS over V2/V3 or over MT/V5, and little or no effect with rTMS over PPC. The results suggest that either the previous psychophysical results with neurological patients are incorrect (highly unlikely) or that the lateral and medial regions are directly connected (as they are in macaque monkeys) such that stimulating one automatically affects the other, in this instance disruptively.

Adult↗

Induced gamma-band oscillations correlate with awareness in hemianopic patient GY.

In normal vision gamma oscillations are involved in object perception, are modulated by attention, and have been linked to awareness by way of their putative role in perceptual integration, arguably as a mechanism for synchronizing activity in separate neural assemblies. We tested the hypothesis that the presence of gamma oscillations (approximately 30-80 Hz) signal the entry of a neural representation into awareness (as indexed by direct report), while attempting to control for other measures of neural information processing such as discrimination accuracy and reaction time. Hemianopic patient GY sometimes reports an awareness "that something happened" in his blind visual hemifield, in response to stimuli of sufficiently high contrast, although he may deny "seeing" anything. At lower contrast levels GY denies any awareness, but may continue to exhibit greater-than-chance accuracy (blindsight). Using a near-threshold level of contrast offers a unique way to test hypotheses concerning correlates of perceptual awareness, since GY's accuracy on certain tasks is independent of awareness. We tested GY on an orientation-discrimination task using stationary stimuli at a fixed near-threshold level of contrast, to which GY sometimes responded "aware" and sometimes "unaware". We recorded brain activity using magnetoencephalography (MEG) in order to determine the relationship between local induced gamma-band oscillations and awareness. GY's accuracy was significantly greater than expected by chance and no different whether or not he reported awareness of the stimulus. Oscillatory activity in the gamma band (44-66 Hz) over the left occipito-parietal region correlated significantly with awareness (but not accuracy), whereas activity in the alpha band (8-12 Hz) did not.

Analysis of Variance↗

Visual area V5/MT remembers "what" but not "where".

Priming for motion direction has been shown to depend upon the functional integrity of extrastriate area V5/MT. Its retinotopic organization and the interactions recently found between motion adaptation and misperceived localization may suggest, for this area, a role for priming of spatial position in addition to the established priming of motion direction. Disruption of V5/MT with repetitive transcranial magnetic stimulation during the intertrial interval had the effect of abolishing priming of motion direction but no effect in priming of spatial position. These effects cannot be explained in terms of perception or task demands but only in terms of the effects of information irrelevant to the correct performance of the task stored over the intertrial interval. We suggest that the attribute of spatial position might be stored in short-term memory either in earlier areas of the motion pathways such as V3 or in higher cortical areas traditionally associated with the analysis of spatial information, for example, posterior parietal cortex or the frontal eye fields.

Adult↗

Do parietal cortical lesions impair spatial attention or allocentric spatial perception?

It remains unclear whether monkeys with large parietal cortical lesions fail "landmark" tasks because they cannot judge the relative distances between landmark and response locations, or because they fail to attend to, or even to notice, the landmark. Monkeys with small posterior parietal (SPP), large posterior parietal (LPP), superior temporal sulcus (STS), or frontal eye field (FEF) lesions were tested on a landmark task in which the physical salience of the landmark and its location varied. Only the LPP monkeys were impaired, seemingly because they overtly failed to shift attention during each trial, responding to whichever food well they looked at first. A task based on one used with neurological patients was therefore introduced in which the monkeys had to discriminate between two white square plaques each containing a spot, where the spot on the positive stimulus was centrally placed. Solving this task requires an allocentric judgement about the relative location of each spot to the edges of the plaque. Even on the most difficult discrimination, monkeys with large parietal lobe lesions were unimpaired. The deficits previously reported on landmark tasks probably reflect a failure of spatial attention or attention to objects rather than an inability to judge allocentric spatial relationships.

Animals↗

The Ferrier Lecture 2004 what can transcranial magnetic stimulation tell us about how the brain works?

Transcranial magnetic stimulation (TMS) is a technique whereby parts of the cerebral cortex and underlying white matter can be excited by a brief electrical current induced by a similarly brief, rapidly fluctuating magnetic field which is itself produced by rapidly discharging a current through an insulated coil held against the scalp. When combined with magnetic resonance structural and functional images of the subject's brain, the stimulation can be directed at specific cortical areas. Over a period of only 15 years, TMS has revealed hitherto unsuspected aspects of brain function, such as the role of distant parts of the brain in recovery from stroke, and has helped to resolve several previously intractable disputes, such as the neuronal basis of conscious awareness. This article describes and discusses the origins and nature of TMS, its applications and limitations, and its especial usefulness in conjunction with other techniques of evaluating or imaging brain activity.

Awareness↗

Deficits of motion integration and segregation in patients with unilateral extrastriate lesions.

Functional neuroimaging in human subjects and single cell recordings in monkeys show that several extra-striate visual areas are activated by visual motion. However, the extent to which different types of motion are processed in different regions remains unclear, although neuropsychological studies of patients with circumscribed lesions hint at regional specialization. We, therefore, studied four patients with unilateral damage to different regions of extrastriate visual cortex on a series of visual discrimination tasks that required them, to a different extent, to integrate local motion signals in order to correctly perceive the direction of global motion. Performance was assessed psychophysically and compared with that of control subjects and with the patients' performance with stimuli presented in the visual field ipsilateral to the lesion. The results indicate considerable regional specialization in extra-striate regions for different aspects of motion processing, namely the largest displacement from frame to frame (D-max) that can sustain perception of coherent motion; perception of relative speed; the amount of coherent motion needed to sustain a percept of global motion in a particular direction; the detection of discontinuities within a moving display; the extraction of form from motion. It was also clear that a defect in local motion, i.e. D-max, can be overcome by integrating local motion signals over a longer period of time. Although no patient suffered from only one defect, the overall pattern of results strongly supports the notion of regional specialization for different aspects of motion processing.

Adult↗

Striate cortex (V1) activity gates awareness of motion.

A key question in understanding visual awareness is whether any single cortical area is indispensable. In a transcranial magnetic stimulation experiment, we show that observers' awareness of activity in extrastriate area V5 depends on the amount of activity in striate cortex (V1). From the timing and pattern of effects, we infer that back-projections from extrastriate cortex influence information content in V1, but it is V1 that determines whether that information reaches awareness.

Awareness↗

Chronostasis without voluntary action.

In a previous study we explored auditory chronostasis and suggested an arousal account of this temporal illusion rather than one dependent on backdating actions to the onset of a motor event. Here we present three experiments designed to distinguish between two competing accounts of the mechanisms underlying the illusion. Experiment 1 investigated whether voluntary movements are necessary for the illusion to occur. Experiment 2 sought to clarify whether auditory chronostasis occurs when the intervals to be judged are continuous (temporally contiguous) rather than separate events. Experiment 3 was designed to establish whether increased task demands account for the illusion. Together the results from these experiments show that chronostasis is an illusion that is not dependent on voluntary action, can occur without a change in the spatial location of the stimulus (thus precluding an account based on spatial attention), occurs with discrete as well as continuous events, and is affected by the salience of the termination of the event to be timed rather than the onset. Collectively these findings suggest that the mechanisms underlying chronostasis are best explained by an arousal hypothesis since neither attention nor backdating to action can account for the commonalities between chronostasis in the auditory, visual and tactile domains.

Acoustic Stimulation↗

Ultrahigh resolution optical coherence tomography of the monkey fovea. Identification of retinal sublayers by correlation with semithin histology sections.

Optical coherence tomography (OCT) has become an established diagnostic tool for the clinical assessment of retinal pathology but correlation of acquired signals with retinal substructures has often been ambiguous. In the monkey retina we have now obtained ultrahigh resolution (UHR) OCT images with 1.4 microm axial x 3 microm transverse resolution from perfusion-fixed eye cups of Macaca fascicularis and optimized the identification of retinal anatomy by correction of spatial artefacts in correlated histology. After resin embedding, serial semithin sections were obtained that corresponded to OCT transects. The direct overlay of features identified in histological sections with corresponding OCT locations was limited by non-linear tissue shrinkage due to dehydration and sectioning stress. In the present study, these misalignments were further corrected by using polygonal spline morphing based on corresponding unequivocal landmarks. The geometric normalization then allowed detailed comparison of both profiles including delicate sublayers of photoreceptor inner- and outer segments. Such correlation will facilitate the extraction of structural information from in vivo ultrahigh resolution OCT images in clinical and experimental applications.

Animals↗

Stimulus cueing in blindsight.

When visual stimuli are presented in the cortically blind visual field of patients or monkeys with verified destruction of striate cortex, many subjects can voluntarily respond to them. In studies of this blindsight, the on- and/or offset of the visual stimulus is usually known to the subject, either because it is signaled in some way or because the subject can present the stimulus himself. To study the effect of stimulus uncertainty on the responses of four hemianopic monkeys and one human hemianope, we compared trials on which the subjects themselves could instantly trigger the stimulus with trials on which the same stimulus appeared 1-7 s after the start-light that normally served as the trigger was first touched. The latter manipulation diminished both the percentage of trials on which the subjects responded and the percentage correct when they did respond. As the start-light disappeared when touched in the first but not second condition, we interpret our results as indicating an influential role for attention in blindsight. Although keeping attention focused on the start-light and delaying the target impaired performance especially in the monkeys, localization was still significant in three and hardly affected in GY.

Animals↗

The 30th Sir Frederick Bartlett lecture. Fact, artefact, and myth about blindsight.

Blindsight is the ability, still controversial if a vote is taken, of subjects with clinically blind field defects to detect, localize, and discriminate visual stimuli of which the subjects say they are completely unaware--the original definition--or of which they might be aware but not in the sense of experiencing a visual percept. These two conditions are known as blindsight Types I and II. This Bartlett lecture narrates the discovery of blindsight and its mounting opposition, and it evaluates the continuing and often perplexing debate about its standing as a visual cognitive phenomenon.

Animals↗

Chromatic priming in hemianopic visual fields.

In three monkeys made hemianopic by unilateral striate cortical ablation, in one normal monkey and in a human hemianope (GY), we measured reaction times to chromatic targets presented in the normal hemifield as a function of prior chromatic primes in the blind field. The first of our three tasks showed an unspecific priming effect in that the colour of the here unpredictive prime was irrelevant. However, when contingencies were changed in the second task so that the prime was usually valid, its colour did significantly influence reaction times in two of the hemianopic monkeys as well as in the human subject. Even when the primes lost their predictive value again in the third task, this chromatically specific effect persisted. We conclude that chromatic processing in the cortically blind field can be revealed with indirect approaches that measure residual processing by its influence on the reaction to stimuli in the normal field, and that the validity of the prime (whether it predicts the colour of the target) is especially important.

Animals↗

Interference with vision by TMS over the occipital pole: a fourth period.

We investigated the effect of single-pulse transcranial magnetic stimulation (TMS) over the occipital pole on a forced-choice visual letter-identification task. Magnetic stimuli were applied on the midline but with the initial current directed pseudorandomly toward either left or right hemisphere; visual stimuli were presented randomly in either left or right hemifield; magnetic-visual stimulus onset asynchrony varied randomly between 12 values: -500 ms and from -50 ms to +50 ms in 10 ms steps. The data revealed the existence of a hitherto unknown fourth task-interfering TMS effect that was maximal at -10 ms and specific for magnetic stimulus polarity and visual stimulus location. This -10 ms effect cannot be explained by reflex blinking (as the -50 ms effect can) and direct disruption of letter-induced activity (as the +20 ms and +100 ms effects can), but it could be explained by direct disruption of pre-letter activity or indirect disruption of letter-induced activity.

Adult↗

The effect of expectation on facilitation of colour/form conjunction tasks by TMS over area V5.

In an earlier paper, we reported task-specific impairments and improvements caused by applying TMS over cortical visual area V5 [Proceedings of the Royal Society of London B 265 (1998) 537]. The phenomenon is further investigated in the present study using two of the previous tasks: a motion/form conjunction in which TMS impaired performance and a colour/form conjunction in which performance was enhanced with TMS. In the earlier experiment, subjects were presented with blocks of trials of one task type perhaps allowing some of the observed effects to arise from knowing the type of stimulus to be discriminated. When blocks of trials consisted of randomly mixed moving/form and colour/form conjunction tasks, TMS over V5 still impaired target-present responses for the moving/form conjunction, but the facilitation seen for colour/form conjunction target-present responses disappeared. We suggest that the competitive inhibition postulated between visual movement areas and colour areas in the brain, in our previous paper, are subject to expectation or knowledge of forthcoming stimulus type.

Adult↗

Can spatial and temporal motion integration compensate for deficits in local motion mechanisms?

We studied the motion perception of a patient, AMG, who had a lesion in the left occipital lobe centered on visual areas V3 and V3A, with involvement of underlying white matter. As shown by a variety of psychophysical tests involving her perception of motion, the patient was impaired at motion discriminations that involved the detection of small displacements of random-dot displays, including local speed discrimination. However, she was unimpaired on tests that required spatial and temporal integration of moving displays, such as motion coherence. The results indicate that she had a specific impairment of the computation of local but not global motion and that she could not integrate motion information across different spatial scales. Such a specific impairment has not been reported before.

Brain Mapping↗