The contribution of human cortical area V3A to the perception of chromatic motion: a transcranial magnetic stimulation study.

Area V3A was identified in five human subjects on both a functional and retinotopic basis using functional magnetic resonance imaging techniques. V3A, along with other visual areas responsive to motion, was then targeted for disruption by repetitive transcranial magnetic stimulation (rTMS) whilst the participants performed a delayed speed matching task. The stimuli used for this task included chromatic, isoluminant motion stimuli that activated either the L-M or S-(L+M) cone-opponent mechanisms, in addition to moving stimuli that contained only luminance contrast (L+M). The speed matching task was performed for chromatic and luminance stimuli that moved at slow (2 degrees/s) or faster (8 degrees/s) speeds. The application of rTMS to area V3A produced a perceived slowing of all chromatic and luminance stimuli at both slow and fast speeds. Similar deficits were found when rTMS was applied to V5/MT+. No deficits in performance were found when areas V3B and V3d were targeted by rTMS. These results provide evidence of a causal link between neural activity in human area V3A and the perception of chromatic isoluminant motion. They establish area V3A, alongside V5/MT+, as a key area in a cortical network that underpins the analysis of not only luminance but also chromatically-defined motion.

Disruptions to human speed perception induced by motion adaptation and transcranial magnetic stimulation.

To investigate the underlying nature of the effects of transcranial magnetic stimulation (TMS) on speed perception, we applied repetitive TMS (rTMS) to human V5/MT+ following adaptation to either fast- (20 deg/s) or slow (4 deg/s)-moving grating stimuli. The adapting stimuli induced changes in the perceived speed of a standard reference stimulus moving at 10 deg/s. In the absence of rTMS, adaptation to the slower stimulus led to an increase in perceived speed of the reference, whilst adaptation to the faster stimulus produced a reduction in perceived speed. These induced changes in speed perception can be modelled by a ratio-taking operation of the outputs of two temporally tuned mechanisms that decay exponentially over time. When rTMS was applied to V5/MT+ following adaptation, the perceived speed of the reference stimulus was reduced, irrespective of whether adaptation had been to the faster- or slower-moving stimulus. The fact that rTMS after adaptation always reduces perceived speed, independent of which temporal mechanism has undergone adaptation, suggests that rTMS does not selectively facilitate activity of adapted neurons but instead leads to suppression of neural function. The results highlight the fact that potentially different effects are generated by TMS on adapted neuronal populations depending upon whether or not they are responding to visual stimuli.

[Abnormal representations in the visual cortex of patients with albinism: diagnostic aid and model for the investigation of the self-organisation of the visual cortex]. / Abnormale Repräsentationen im visuellen Kortex von Albinismus-Patienten : Hilfsmittel bei der Diagnostik und Modell der kortikalen Selbstorganisation.

A characteristic feature of patients with albinism is the misrouting of the optic nerves, which causes the visual cortex to receive an abnormal input. This report details how the detection of misrouting using visual evoked potentials assists the clinical diagnosis of albinism. Further, it shows how the projection abnormality observed in patients with albinism provides a model for investigating the self-organisation of the human visual cortex. This is highlighted by recent findings that were obtained using functional magnetic resonance imaging, visual evoked potentials, and static visual field perimetry.

Abnormal visual projection in a human albino studied with functional magnetic resonance imaging and visual evoked potentials.

The albino visual pathway is abnormal in that many fibres from the temporal retina project to the contralateral visual cortex. The visual projections in a human albino and a control have been investigated with fMRI and VEP during independent visual stimulation of both hemifields. Activity in the occipital cortex in the normal was contralateral to the stimulated visual field, whereas it was contralateral to the stimulated eye in the albino, independent of the stimulated visual field. Thus, the albino visual cortex is activated not only by stimulation in the contralateral visual field, but also by abnormal input representing the ipsilateral visual field. These novel findings help elucidate the nature of albino misrouting.

Abnormal retinotopic representations in human visual cortex revealed by fMRI.

The representation of the visual field in early visual areas is retinotopic. The point-to-point relationship on the retina is therefore maintained on the convoluted cortical surface. Functional magnetic resonance imaging (fMRI) has been able to demonstrate the retinotopic representation of the visual field in occipital cortex of normal subjects. Furthermore, visual areas that are retinotopic can be identified on computationally flattened cortical maps on the basis of positions of the vertical and horizontal meridians. Here, we investigate abnormal retinotopic representations in human visual cortex with fMRI. We present three case studies in which patients with visual disorders are investigated. We have tested a subject who only possesses operating rod photoreceptors. We find in this case that the cortex undergoes a remapping whereby regions that would normally represent central field locations now map more peripheral positions in the visual field: In a human albino we also find abnormal visual cortical activity. Monocular stimulation of each hemifield resulted in activations in the hemisphere contralateral to the stimulated eye. This is consistent with abnormal decussation at the optic chiasm in albinism. Finally, we report a case where a lesion to white matter has resulted in a lack of measurable activity in occipital cortex. The activity was absent for a small region of the visual field, which was found to correspond to the subject's field defect. The cases selected have been chosen to demonstrate the power of fMRI in identifying abnormalities in the cortical representations of the visual field in patients with visual dysfunction. Furthermore, the experiments are able to show how the cortex is capable of modifying the visual field representation in response to abnormal input.

Diurnal daylight phase affects the temporal properties of both the b-wave and d-wave of the human electroretinogram.

Aspects of the anatomy and physiology of the cone pathway are known to vary according to the phase of the natural light cycle. Using a prolonged flash stimulus ( approximately 200 ms), we have examined the human electroretinogram (ERG) over a 24 h period. We report that whilst the a-wave of the photopic ERG does not alter, there are profound effects upon the implicit times of both the b-wave and d-wave components. Both components are significantly slower in the night-time period and systematically become faster (15-22% reduction in implicit time), reaching a peak at around midday. The daily variation in the temporal properties of the ERG is abolished by constant light, but is retained during constant darkness. The data suggest that the changes in the temporal properties of the cone pathway affect both cone-ON and cone-OFF pathways. This suggests that the diurnal effect is presynaptic to the second order neurones, and most likely resides in the cone synapse.

Scanning the visual world: a study of patients with homonymous hemianopia.

OBJECTIVES: This study examined the scanpaths of patients with homonymous hemianopia while viewing naturalistic pictures in their original and also spatially filtered forms. Features of their scanpaths with respect to various saccade and fixation parameters were examined to determine whether they develop compensatory eye movement strategies. The effects of various lesion parameters including location, size, and age on the evolution of such strategies were considered. METHODS: Eye movements of eight patients with homonymous hemianopia (four left, four right), but lacking neglect, were recorded while they viewed 22 images of real scenes, and they were compared with the eye movements of eight age matched controls. Subjects viewed each image for 3 seconds, initially in a spatially filtered form in which much of the semantic content had been removed, and then in their unfiltered, original form. RESULTS: Patients differed significantly from controls in various fixation and saccade parameters. For fixation parameters patients with hemianopia fixated different spatial positions from controls, made more fixations which were more widely distributed and of shorter duration than controls, and spent a greater proportion of their total fixation time in the area corresponding to their blind hemifield. They did not make significantly more refixations than controls. For saccade parameters patients made more saccades into their blind hemifield, these saccades having shorter latencies and shorter amplitudes than those made into their seeing field, and had longer scanpaths than control subjects. The amplitude of their first saccade was longer than that of controls although its direction did not correlate simply with the side of the field defect. Their mean saccade amplitude was similar to that of controls. Filtering out high spatial frequencies within images seemed to accentuate the described differences between eye movement characteristics of hemianopes and controls. Scanpath differences correlated with increasing age but not location or size of lesions causing the hemianopia. CONCLUSION: Various features of scanpaths produced by hemianopes were different from normal subjects. These differences correlated with lesion age and may reflect the evolution of a compensatory eye movement strategy.

Adaptation to oscillopsia: a psychophysical and questionnaire investigation.

In this study we explore the reasons why patients with bilateral vestibular failure report disparate degrees of oscillopsia. Twelve bilateral labyrinthine-defective (LD) subjects and twelve normal healthy controls were tested using a self- versus visual-motion psychophysical experiment. The LD subjects also completed a questionnaire designed to quantify the severity of handicap caused by oscillopsia. Additional standardized questionnaires were completed to identify the role of personality, personal beliefs and affective factors in adaptation to oscillopsia. During the psychophysical experiment subjects sat on a motorized Barany chair whilst viewing a large-field projected video image displayed on a screen in front of them. The chair and video image oscillated sinusoidally at 1 Hz in counter-phase at variable amplitudes which were controlled by the subject but constrained, so that the net relative motion of the chair and video image always resulted in a sinusoid with a peak velocity of 50 degrees /s. The subject's task was to find the ratio of chair versus video image motion that subjectively produced the 'most comfortable visual image'. Eye movements were recorded during the experiment in order that the net retinal image slip at the point of maximum visual comfort could be measured. The main findings in the LD subjects were that, as a group, they selected lower chair motion amplitude settings to obtain visual comfort than did the normal control subjects. Responses to the questionnaires highlighted considerable variation in reported handicap due to oscillopsia. Greater oscillopsia handicap scores were significantly correlated with a greater external locus of control (i.e. the perception of having little control over one's health). Retinal slip speed was negatively correlated with oscillopsia handicap score so that patients who suffered the greatest retinal slip were those least handicapped by oscillopsia. The results suggest that adaptation to oscillopsia is partly related to the patient's personal attitude to the recovery process and partly associated with the development of tolerance to the movement of images on the retina during self-motion. The latter is likely to be related to previously described changes in visual motion sensitivity in these patients.

Visual perception of motion, luminance and colour in a human hemianope.

Human patients rendered cortically blind by lesions to V1 can nevertheless discriminate between visual stimuli presented to their blind fields. Experimental evidence suggests that two response modes are involved. Patients are either unaware or aware of the visual stimuli, which they are able to discriminate. However, under both conditions patients insist that they do not see. We investigate the fundamental difference between percepts derived for the normal and affected hemifield in a human hemianope with visual stimuli of which he was aware. The psychophysical experiments we employed required the patient, GY, to make comparisons between stimuli presented in his affected and normal hemifields. The subject discriminated between, and was allowed to match, the stimuli. Our study reveals that the stimulus parameters of colour and motion can be discriminated and matched between the normal and blind hemifields, whereas brightness cannot. We provide evidence for associations between the percepts of colour and motion, but a dissociation between the percepts of brightness, derived from the normal and hemianopic fields. Our results are consistent with the proposal that the perception of different stimulus attributes is expressed in activity of functionally segregated visual areas of the brain. We also believe our results explain the patient's insistence that he does not see stimuli, but can discriminate between them with awareness.

Topographic organization of human visual areas in the absence of input from primary cortex.

Recently, there has been evidence for considerable plasticity in primary sensory areas of adult cortex. In this study, we asked to what extent topographical maps in human extrastriate areas reorganize after damage to a portion of primary visual (striate) cortex, V1. Functional magnetic resonance imaging signals were measured in a subject (G.Y.) with a large calcarine lesion that includes most of primary visual cortex but spares the foveal representation. When foveal stimulation was present, intact cortex in the lesioned occipital lobe exhibited conventional retinotopic organization. Several visual areas could be identified (V1, V2, V3, V3 accessory, and V4 ventral). However, when stimuli were restricted to the blind portion of the visual field, responses were found primarily in dorsal extrastriate areas. Furthermore, cortex that had formerly shown normal topography now represented only the visual field around the lower vertical meridian. Several possible sources for this reorganized activity are considered, including transcallosal connections, direct subcortical projections to extrastriate cortex, and residual inputs from V1 near the margin of the lesion. A scheme is described to explain how the reorganized signals could occur based on changes in the local neural connections.

Oscillopsia: visual function during motion in the absence of vestibulo-ocular reflex.

OBJECTIVES: To investigate (1) the effects of loss of vestibular function on spatiotemporal vision and (2) the mechanisms which enable labyrinthine defective (LD) patients to adapt to oscillopsia. METHODS: Visual function and eye movements were assessed in seven normal subjects and four LD patients with oscillopsia due to absent vestibulo-ocular reflex. Temporal vision was assessed by measurement of threshold sensitivity for detection of a target which moved across a flickering, spatially uniform background field. Spatial vision was investigated by measurements of threshold sensitivity for the detection of a target moving across a spatially modulated background in the form of square wave gratings. Velocity discrimination was assessed with drifting gratings. All measurements were made under static conditions and during oscillatory movement of either the visual stimulus or the subject (1 Hz, peak velocity 50 degrees/s). RESULTS: Temporal responses--Normal subjects and LD patients exhibited similar responses while static and under body oscillation. Spatial responses--The two groups achieved similar results under static conditions but body oscillation reduced threshold sensitivities and shifted the spatial response function towards lower spatial frequencies in the LD patients only. Similar changes in the spatial responses were seen during oscillation of the visual stimulus but these occurred in both normal subjects and LD patients. Velocity discrimination--Two LD patients achieved normal velocity discrimination but the other two showed abnormal responses to visual stimulus movement; one displayed a loss of velocity discrimination during whole body oscillation, and the other mismatched the velocity of two moving grating stimuli. CONCLUSIONS: The changes in the spatial responses are attributed to the presence of retinal slip during visual stimulus motion in all subjects or body oscillation in the LD patients. It is concluded that any visual adaptation to oscillopsia achieved by the LD patients does not influence the measured spatial response functions, which arise at an early stage of visual processing. The abnormal velocity discrimination may relate to the progressive improvement in oscillopsia reported by LD patients.

Movement in the normal visual hemifield induces a percept in the 'blind' hemifield of a human hemianope.

We have investigated visual responses to moving stimuli presented to the normal hemifield of a hemianope, GY, who exhibits residual visual function in his right, 'blind' hemifield. Preliminary experiments established that his perception of moving stimuli localized in his 'blind' hemifield is retained when a similar stimulus is presented simultaneously in the normal hemifield. In response to a grating stimulus moving horizontally towards fixation in the non-foveal region of the normal, left hemifield, he perceives in addition to a normal motion percept in the left hemifield, a sensation of movement localized in the right hemifield. Qualitatively, this latter is indistinguishable from responses elicited by direct stimulation localized within his 'blind' hemifield by moving stimuli. We have investigated the characteristics of the mechanisms which induce the 'blind' field component of GY's responses to stimulation of the normal hemifield. We show that GY's sensitivity for detection of movement localized within his 'blind' hemifield is dependent on the direction of movement, the contrast and the velocity of a grating presented to the normal hemifield. No induced effects were recorded in response to colour or to non-moving, flickering stimuli. We examine the possible contribution of scattered light to our observations, and eliminate this factor by consideration of our experimental results. We discuss the neural mechanisms which may be involved in this response.

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.

A new abnormality of human vision provides evidence of interactions between cortical mechanisms sensitive to movement and those sensitive to colour.

We present the results of a psychophysical study on a human observer, MW, which describe his abnormal visual responses to moving stimuli. It has been shown previously that this subject has normal vision for stationary, achromatic patterns, but responds highly abnormally to saturated chromatic, and especially red stimuli. We now report that like chromatic stimuli, moving black and white patterns elicit an inhibitory response which extends beyond the visual field area covered by the moving stimulus itself, and suppresses detection of stationary, achromatic patterns. Although both chromatic and moving stimuli generate similar percepts in association with their inhibitory activities, these latter differ in several respects, and we conclude that they have different neural origins. We show that in addition to their separate inhibitory actions on detection of achromatic patterns, movement and colour exert mutually inhibitory effects. Movement is markedly effective in limiting the inhibitory spread associated with colour, regardless of the positions in the visual field of the moving and coloured stimuli. Such spatially diffuse activity is characteristic of higher visual processing in pre-occipital cortical visual areas.

Orientation discrimination is impaired in the absence of the striate cortical contribution to human vision.

We have examined visual discrimination of orientation for bars and edges presented to the 'blind' hemifield of a patient, GY, who has an extensive lesion involving the left striate and prestriate cortex. The patient's residual vision in response to transient light stimulation of his 'blind' hemifield has been well characterized, and it provides discrimination on the basis of differences in target velocity, flicker frequency or colour. We now show that using his residual vision, GY can identify accurately the orientation of a flickering bar, although his performance is sub-normal for bars shorter than 10 deg. He is, however, unable to identify the orientation of a one-dimensional grating presented within a circular aperture, and is unable to detect a grating structure formed by equiluminant coloured bars or by alternating, equiluminant flickering and steady bars. We also show that he has poor orientation discrimination for edges formed by colour contrast, by contrast between moving stimuli or by flicker contrast. We conclude that in the absence of the striate cortical input, the residual mechanisms which provide discriminations for colour, flicker or movement are not well organized for the detection and discrimination of stimulus orientation. We discuss the physiological mechanisms responsible for the various aspects of GY's residual vision.

Object motion perception during ego-motion: patients with a complete loss of vestibular function vs. normals.

Object motion perception was assessed in avestibular patients and normal controls. Two experiments were conducted, in which subjects were required to assess the motion of a visual stimulus with respect to earth. In the first experiment, we measured the velocity at which a briefly presented (200 ms) grating was perceived as earth fixed, while the subject maintained fixation on a visual target fixed relative to the body, during whole-body yaw rotation (VOR suppression). In this experimental setup, the influence of the semicircular canal signals on object motion perception was evaluated. The avestibular patients judged the grating to be stationary with respect to earth, when it was moving at the same velocity as their body, whereas for normal controls, the grating was perceived as stationary when it moved at a velocity slower than their body motion, but greater than zero. The difference between the two subject groups was significant, and showed the strong contribution of the vestibular system to object motion perception. Similarly, a measurement of the velocity at which a grating was perceived as stationary was obtained during smooth pursuit eye movements. In this experiment the contribution of the efference copy of the oculomotor signal and proprioceptive signals to object motion perception were assessed. As with the first experiment, the normal controls displayed a more veridical sense of object motion perception than the patients, although the difference was only just significant. We suggest that the difference could be an adaptive change in the patients perception of motion, which allows them to reduce the effects of oscillopsia.

Colour identification and colour constancy are impaired in a patient with incomplete achromatopsia associated with prestriate cortical lesions.

We have examined visual functions, including colour vision, in a patient with bilateral cortical lesions involving mainly the fusiform and lingual gyri, areas known to be involved in the central processing of chromatic stimuli. The patient has near normal (6/9) acuity, and his responses to tests of binocular function and spatial vision are normal, as are his discrimination of changes in target speed and surface lightness. He does, however, exhibit minor losses in the upper visual field, mild prosopagnosia and topographical agnosia, all conditions commonly associated with cerebral achromatopsia. Colour matches and spectral response data establish that his cone photoreceptors have normal spectral characteristics and his spectral sensitivity measured against a white background reveals normal postreceptoral chromatic function. The patient's colour discrimination for differences in wavelength, hue or saturation is, however, impaired and his colour naming is significantly disturbed, particularly for blues and greens. We have determined the areas of the chromaticity chart that correspond to his naming categories for surface colours, and show that changes in illuminant cause him to alter the names of surface colours in a manner consistent with the changes in their chromaticities. Other subjects with normal or congenital red-green deficient colour vision make many fewer name changes under changes in illuminant. We conclude that the patient's colour constancy is impaired as a consequence of abnormal central processing of colour vision.

Vision during motion in patients with absent vestibular function.

We have measured a spatial visual response and visual velocity discrimination in 4 patients with long standing vestibular loss and 6 controls. The spatial response was measured during; i) body and visual display stationary conditions, ii) whole-body oscillation (1 Hz +/- 50 degrees/s) and iii) visual stimulus oscillation (1 Hz +/- 50 degrees/s). Velocity discrimination was assessed during conditions i) and ii). The visual tests applied were selected on the basis that the spatial response is known to reflect peripheral processes of the retina, whereas velocity processing is more central in origin. Patients had normal spatial responses under static conditions and they suffered a degradation in their spatial responses during whole-body oscillation, whereas, normals' responses remained unaltered. During oscillation of the visual display both patients and normals suffered a degradation in their spatial responses, and for patients the change was very similar to that observed during whole-body oscillation. The changes in the spatial responses were dependent on the gain of the eye movements which compensated for the whole-body or visual display oscillation. In 3 patients and all controls whole-body oscillation did not alter the discrimination of velocity of a vertically moving horizontally orientated grating compared with when the subjects were stationary. One patient suffered a severe reduction in the ability to discriminate velocity under whole-body oscillation, which suggests that central suppression of motion perception reduces oscillopsia.

Recovery from bilateral vestibular failure: implications for visual and cervico-ocular function.

We report a patient who sustained severe bilateral labyrinthine lesions during Streptococcus suis meningitis but considerably recovered vestibular function over a 7 month period. This unique case allowed us to examine the cervico-ocular reflex (COR) and visual function at various levels of activity of his vestibular system. The slow phase COR, elicited by trunk oscillation (0.2 Hz) with the head earth-stationary, was negligible immediately after the acute vestibular loss but rose to a gain of 0.51 one month after. Seven months later, when vestibular function was improved, COR gain dropped to a gain of 0.15. Measurements of spatial visual function during whole body oscillation in the acute stage and after 6 months showed marked improvement which correlated entirely with VOR measurements in the dark and during optic fixation. This patient also showed the unique feature that, in the acute stage, eye movement gain and visual function were poorer during whole body motion than during identical visual target motion. These findings suggest that: i) the COR may be inhibited by the presence of vestibular signals, ii) spatial vision measurements provide accurate assessment of the patient's visual blur during head motion, and iii) the severe oscillopsia experienced by patients in the acute stage of vestibular loss may not only be due to the absence of the VOR; additional degradation in eye movements during head motion, perhaps arising from acutely distorted labyrinthine signals, may also play a part.

Spatial, colour and contrast response characteristics of mechanisms which mediate discrimination of pattern orientation and magnification.

We have measured response times for the detection of a single target presented against a set of reference elements which are characterised by combinations of four different stimulus parameters; colour, contrast polarity, magnification and orientation. The aim of the experiments was to determine the response characteristics of visual mechanisms which mediate target detection through the discrimination of orientation and magnification. In the first experiments, we determined sensitivity to differences in colour and contrast polarity, and show that the mechanisms responsible for the discrimination of orientation and of magnification are both selective in their responses to colour and to contrast polarity. There are, nonetheless, residual interactions between patterns of different contrast polarities and between those of different colour, and in the latter case, weak interactions persist under equiluminance conditions. In a second set of experiments, we examined the interactions between orientation and magnification. We conclude that the responses of visual mechanisms which mediate target detection through discrimination of orientation are markedly dependent on stimulus magnification whereas those which mediate detection through discrimination of magnification are, in contrast, relatively insensitive to stimulus orientation.