A motor signal and “visual” size perception

Recent models of the visual system in primates suggest that the mechanisms underlying visual perception and visuomotor control are implemented in separate functional streams in the cerebral cortex. However, a little-studied perceptual illusion demonstrates that a motor-related signal representing arm position can contribute to the visual perception of size. The illusion consists of an illusory size change in an afterimage of the hand when the hand is moved towards or away from the subject. The motor signal necessary for the illusion could be specified by feedforward and/or feedback sources (i.e. efference copy and/or proprioception/kinesthesis). We investigated the nature of this signal by measuring the illusion's magnitude when subjects moved their own arm (active condition, feedforward and feedback information available), and when arm movement was under the control of the experimenter (passive condition, feedback information available). Active and passive movements produced equivalent illusory size changes in the afterimages. However, the illusion was not obtained when an afterimage of subject's hand was obtained prior to movement of the other hand from a very similar location in space. This evidence shows that proprioceptive/kinesthetic feedback was sufficient to drive the illusion and suggests that a specific three-dimensional registration of proprioceptive input and the initial afterimage is necessary for the illusion to occur.     

Temporal recalibration during asynchronous audiovisual speech perception

When a participant moves a hand-held target in complete darkness after an afterimage of that target has been obtained, an illusory increase (with movements away from the participant) or decrease (with movements towards the participant) in the apparent size of the afterimage is reported (the Taylor illusion, reported first in Taylor, J Exp Psychol 29: 1941). Unlike typical Emmert’s Law demonstrations, the Taylor illusion shows that a motor-related signal can be used to specify distance for the computation of real size. A study by Carey and Allan (Exp Brain Res 110: 1996) found that the Taylor illusion did not occur in a condition where an afterimage of one hand was obtained while the other hand performed a movement away from the participant from directly behind the first. It was proposed that, for the illusion to manifest itself, proprioceptive and visual information must be in strict “register” when the afterimage is obtained. To evaluate this hypothesis, 14 participants performed “towards” and “away” movements after obtaining afterimages of hand-held cards. Participants wore either plain lenses or prism lenses during the trials, the latter of which displaced visual stimuli 10° to the left. No significant difference was found between the two lens conditions in terms of the effect on the perception of the Taylor illusion. It was concluded that the illusory size distortions may depend on register of visual and proprioceptive position in terms of depth, rather than in the picture plane. Several suggestions for future studies of the Taylor illusion are proposed, and limitations of size judgements of afterimages are outlined.     

Suppression of OKN and VOR by afterimages and imaginary objects

Summary Optokinetic nystagmus (OKN) is suppressed if attention is directed to a centrally placed afterimage superimposed on a moving display. Imagining a stationary object has little or no effect. An afterimage does not provide the retinal slip and misfoveation error signals provided by a stationary object and we have shown that an effective error signal does not arise from occlusion or masking of the display by the afterimage. Although a lack of relative motion between afterimage and moving display could indicate when OKN gain is one, there is no unique relative motion signal associated with a gain of zero. Subjects could partially inhibit the vestibulo-ocular reflex (VOR) in the dark when they imagined a head-fixed object. They could suppress the response more effectively by attending to an afterimage, but the suppression was still only partial. When OKN and VOR were evoked simultaneously, pursuit movements of the eyes could not be suppressed until the vestibular inputs had subsided. We conclude that signals associated with OKN, are fully available to the mechanism that assesses the headcentric motion of objects but that signals associated with VOR are only partially available to that mechanism.This study was part of DCIEM research contract 97711-3-7595/ 8SE83-00221     

Looking at the task in hand: vergence eye movements and perceived size

A retinal afterimage of the hand changes size when the same unseen hand is moved backwards and forwards in darkness. We demonstrate that arm movements per se are not sufficient to cause a size change and that vergence eye movements are a necessary and sufficient condition for the presence of the illusory size change. We review previous literature to illustrate that changing limb position in the dark alters vergence angle and we explain the illusion via this mechanism. A discussion is provided on why altering limb position causes a change in vergence and we speculate on the underlying mechanisms. Received: 6 May 1997 / Accepted: 16 July 1997     

Predicting the orientation of invisible stimuli from activity in human primary visual cortex

Humans can experience aftereffects from oriented stimuli that are not consciously perceived, suggesting that such stimuli receive cortical processing. Determining the physiological substrate of such effects has proven elusive owing to the low spatial resolution of conventional human neuroimaging techniques compared to the size of orientation columns in visual cortex. Here we show that even at conventional resolutions it is possible to use fMRI to obtain a direct measure of orientation-selective processing in V1. We found that many parts of V1 show subtle but reproducible biases to oriented stimuli, and that we could accumulate this information across the whole of V1 using multivariate pattern recognition. Using this information, we could then successfully predict which one of two oriented stimuli a participant was viewing, even when masking rendered that stimulus invisible. Our findings show that conventional fMRI can be used to reveal feature-selective processing in human cortex, even for invisible stimuli.     

Discriminating the role of binocular information in the timing of a one-handed catch

The effects of early, unilateral cerebral hemispherectomy on retinogeniculate projections were studied in the vervet monkey (Cercopithecus aethiops sabeus).Hemispherectomy eliminates all geniculocortical pathways and thus removes cortical factors involved in the survival of retinogeniculate projections. Complete removal of the left cerebral cortex was performed in two monkeys at 6 months and 8 months of age. After a postsurgical survival period of 50 months (SHG3) and 45 months (SHG4), both animals and a normal adult monkey received intraocular injections of [³H]proline (5 mCi) in the left eye and WGA-HRP (100 μl, 5%) in the right eye. The dorsal lateral geniculate nucleuseuron (LGNd) ipsilateral to the hemispherectomy was on average 73% smaller than the contralateral LGNd. The magno- and parvocellular layers ipsilateral to the cortical ablation in both hemispherectomized subjects received a layered, eye-specific pattern of retinal input. This suggests that retinogeniculate projections could be sustained in the absence of geniculate relay cells. Electronic Publication     

EEG Correlates of the Afterimage of Visual Stimulation

Occipital EEG alpha activity following visual stimuli of varying intensity was studied with particular reference to the visual afterimage. Subjects each received three stimuli of differing intensity under two conditions: with and without reporting of afterimage onset and offset required. A third condition controlled for possible reporting effects on the EEG. Findings indicate that visual afterimages are a significant determinant of the duration of EEG alpha desynchronization following visual stimulation and that where visual stimuli are employed the duration of EEG alpha blocking is unsuitable as an indicator of the OR to such stimulation.     

Cerebellar lesions impair context-dependent adaptation of reaching movements in primates

To produce accurate movements when conditions change suddenly, the brain must be capable of learning multiple versions of a given motor task and must be able to access the appropriate program using sensory information linked to the context of the movement. The neural basis for context-dependent motor learning is uncertain, but the cerebellum is thought to play a fundamental role. In this study, we examined the effect of lesions of the dorsal vermal and paravermal cerebellar cortex on the adaptation of reaching movements produced by modified visual feedback and accessed with a visual cue. Two rhesus monkeys were trained to point to targets displayed on a video monitor while viewing monocularly with either eye. During the experimental sessions, visual information received by one eye (the "modified" eye) was displaced horizontally, while the information received by the other ("normal") eye remained unaltered. In the first set of experiments (noncontextual paradigm), the animals pointed to targets while viewing with the modified eye. This paradigm resulted in a gradual improvement in pointing accuracy when viewing with that eye, but also produced a shift in pointing responses of equivalent size when viewing with the normal eye. In the second set of experiments (contextual paradigm), the animals alternated six blocks of reaches while viewing monocularly with the modified eye with six blocks viewing with the normal eye. This paradigm improved the pointing accuracy when viewing with the modified eye, but produced only a small shift in pointing responses when viewing with the normal eye. After the dorsal vermal and paravermal cerebellar cortex were resected, no change occurred in the pattern of adaptation produced by the noncontextual paradigm. The contextual paradigm, however, no longer selectively adapted pointing responses for each eye, but rather produced a pointing shift of equivalent size when viewing with either eye. The results indicate that pointing responses can be differentially adapted for each viewing eye, which is a form of context-dependent motor learning. This capability was lost after focal lesions of the dorsal vermal and paravermal cerebellar cortex, suggesting that these regions of cerebellar cortex are required to learn or store multiple representations of a movement, or to retrieve the appropriate motor program in a given sensory context.     

Adaptive modification of saccade size produces correlated changes in the discharges of fastigial nucleus neurons

Saccade accuracy is known to be maintained by adaptive mechanisms that progressively reduce any visual error that consistently exists when the saccade ends. We used an experimental paradigm known to induce adaptation of saccade size while monitoring the neural correlates of this adaptation. In rhesus monkeys where the medial and lateral recti of one eye were surgically weakened, patching the unoperated eye and forcing the monkey to use the weakened eye induced a gradual increase in saccade size in both eyes until the viewing, weak eye almost acquired the target in one step. Subsequent patching of the weakened eye gradually reversed the situation, so that the saccades in the viewing, normal eye decreased from an initial overshooting to normal. In the caudal fastigial nuclei of unadapted monkeys, neurons typically exhibit an early burst of spikes that is correlated with the onset of contraversive saccades and a later burst of spikes that is correlated with the termination of ipsiversive saccades. Comparing the discharges of the same fastigial neurons recorded before and during adaptation, this basic pattern did not change, but some parameters of the discharges did. The most consistent changes were in the latency of the burst for ipsiversive saccades, which was positively correlated with saccade size (1.28 ms/deg), and in the number of spikes associated with contraversive saccades, which was also positively correlated (0.55 spikes/deg). The former was more important when saccade size was decreasing, and the latter was more important when saccade size was increasing. Based on current knowledge of the anatomical connections of fastigial neurons, as well as on the effects of cerebellar lesions and on recordings in other structures, we argue that these changes are appropriate for causing the associated changes in saccade size.     

Analyses of responsible factors for the delay effect produced by prolonged viewing in recognizing Kanji characters

Ninose & Gyoba (1996) examined delays in the recognition of Kanjis following prolonged viewing. They suggested that a Kanji pattern may be internally represented as a whole and the prolonged viewing may produce an adaptation effect specific to such a representation. The present study examined responsible factors producing the delay effect more in detail by manipulating figural components of adaptation and test Kanjis. The delay occurred only when the test Kanji was the same pattern as the adaptation Kanji in all local details. In contrast, the delay were not observed either when the global pattern of the test Kanji was similar to that of the adaptation Kanji, or when a part of the test Kanji was the same pattern as the whole configuration of the adaptation Kanji. These results indicate that the delay effect is produced by prolonged viewing of the adaptation Kanji with exactly the same local and global configurations as the test Kanji, while the previous studies have revealed that the delay effect occurs independent from Kanji's size and orientation in some ranges.     

Human ocular responses to translation of the observer and of the scene: dependence on viewing distance

Recent experiments on monkeys have indicated that the eye movements induced by brief translation of either the observer or the visual scene are a linear function of the inverse of the viewing distance. For the movements of the observer, the room was dark and responses were attributed to a translational vestibulo-ocular reflex (TVOR) that senses the motion through the otolith organs; for the movements of the scene, which elicit ocular following, the scene was projected and adjusted in size and speed so that the retinal stimulation was the same at all distances. The shared dependence on viewing distance was consistent with the hypothesis that the TVOR and ocular following are synergistic and share central pathways. The present experiments looked for such dependencies on viewing distance in human subjects. When briefly accelerated along the interaural axis in the dark, human subjects generated compensatory eye movements that were also a linear function of the inverse of the viewing distance to a previously fixated target. These responses, which were attributed to the TVOR, were somewhat weaker than those previously recorded from monkeys using similar methods. When human subjects faced a tangent screen onto which patterned images were projected, brief motion of those images evoked ocular following responses that showed statistically significant dependence on viewing distance only with low-speed stimuli (10°/s). This dependence was at best weak and in the reverse direction of that seen with the TVOR, i.e., responses increased as viewing distance increased. We suggest that in generating an internal estimate of viewing distance subjects may have used a confounding cue in the ocular-following paradigm the size of the projected scene -which was varied directly with the viewing distance in these experiments (in order to preserve the size of the retinal image). When movements of the subject were randomly interleaved with the movements of the scene to encourage the expectation of ego-motion -the dependence of ocular following on viewing distance altered significantly: with higher speed stimuli (40°/s) many responses (63%) now increased significantly as viewing distance decreased, though less vigorously than the TVOR. We suggest that the expectation of motion results in the subject placing greater weight on cues such as vergence and accommodation that provide veridical distance information in our experimental situation: cue selection is context specific.     

Human ocular responses to translation of the observer and of the scene: dependence on viewing distance

Recent experiments on monkeys have indithat-—the eye movements induced by brief translation of either the observer or the visual scene are a linear function of the inverse of the viewing distance. For the movements of the observer, the room was dark and responses were attributed to a translational vestibulo-ocular reflex (TVOR) that senses the motion through the otolith organs; for the movements of the scene, which elicit ocular following, the scene was projected and adjusted in size and speed so that the retinal stimulation was the same at all distances. The shared dependence on viewing distance was consistent with the hypothesis that the TVOR and ocular following are synergistic and share central pathways. The present experiments looked for such dependencies on viewing distance in human subjects. When briefly accelerated along the interaural axis in the dark, human subjects generated compensatory eye movements that were also a linear function of the inverse of the viewing distance to a previously fixated target. These responses, which were attributed to the TVOR, were somewhat weaker than those previously recorded from monkeys using similar methods. When human subjects faced a tangent screen onto which patterned images were projected, brief motion of those images evoked ocular following responses that showed statistically significant dependence on viewing distance only with low-speed stimuli (10°/s). This dependence was at best weak and in the reverse direction of that seen with the TVOR, i.e., responses increased as viewing distance increased. We suggest that in generating an internal estimate of viewing distance subjects may have used a confounding cue in the ocular-following paradigmthe size of the projected scene - which was varied directly with the viewing distance in these experiments (in order to preserve the size of the retinal image). When movements of the subject were randomly interleaved with the movements of the scene - to encourage the expectation of ego-motion - the dependence of ocular following on viewing distance altered significantly: with higher speed stimuli (40°/s) many responses (63%) now increased significantly as viewing distance decreased, though less vigorously than the TVOR. We suggest that the expectation of motion results in the subject placing greater weight on cues such as vergence and accommodation that provide veridical distance information in our experimental situation: cue selection is context specific.     

Viewing the body modulates tactile receptive fields

Tactile discrimination performance depends on the receptive field (RF) size of somatosensory cortical (SI) neurons. Psychophysical masking effects can reveal the RF of an idealized “virtual” somatosensory neuron. Previous studies show that top–down factors strongly affect tactile discrimination performance. Here, we show that non-informative vision of the touched body part influences tactile discrimination by modulating tactile RFs. Ten subjects performed spatial discrimination between touch locations on the forearm. Performance was improved when subjects saw their forearm compared to viewing a neutral object in the same location. The extent of visual information was relevant, since restricted view of the forearm did not have this enhancing effect. Vibrotactile maskers were placed symmetrically on either side of the tactile target locations, at two different distances. Overall, masking significantly impaired discrimination performance, but the spatial gradient of masking depended on what subjects viewed. Viewing the body reduced the effect of distant maskers, but enhanced the effect of close maskers, as compared to viewing a neutral object. We propose that viewing the body improves functional touch by sharpening tactile RFs in an early somatosensory map. Top–down modulation of lateral inhibition could underlie these effects.     

Ocular responses to translation and their dependence on viewing distance. II. Motion of the scene.

1. The ocular following responses induced by brief (100-ms) movements of the visual scene were examined for their dependence on viewing distance in 5 monkeys (Macaca mulatta). The horizontal positions of both eyes and the vertical position of one eye were recorded using the electromagnetic search-coil technique. Accommodation was monitored in selected experiments by use of an infrared optometer. Test patterns (random dots) were back-projected onto a translucent tangent screen facing the animal. Six viewing distances were used (range, 20-150 cm), the size and speed of the image on the screen being adjusted for each so as to preserve a constant retinal image. 2. Response measures based on the amplitude of the first peak in the eye acceleration profile or the eye velocity achieved at specific times all indicated that ocular following responses were inversely related to viewing distance, the relationship being linear for the earliest measures. On average, the sensitivity to viewing distance was comparable with that reported for the translational vestibuloocular reflex (TVOR) in the preceding paper: as viewing distance increased from 20 cm, ocular following decremented at a mean rate (+/- SD) of 17 +/- 3% per m-1, while the TVOR decremented at a mean rate (+/- SD) of 18 +/- 1% per m-1. 3. Ocular following responses showed the postsaccadic enhancement described by Kawano and Miles regardless of viewing distance. To a first approximation, the effects of postsaccadic enhancement and viewing distance summed linearly. 4. The dependence of ocular following on speed showed the progressive saturation previously described by Miles et al. at all viewing distances, the peak eye velocity achieved being inversely related to the viewing distance, indicating that the saturation must originate upstream of the dependence on viewing distance. Under normal viewing conditions, this speed saturation will tend to offset the dependence on viewing distance because the retinal slip speeds experienced by the moving observer will tend to vary inversely with viewing distance, resulting in greater saturation with nearer viewing. 5. Wedge prisms were used to dissociate vergence and accommodation and indicated that ocular following responses were sensitive to selective increases in either vergence (base-out prism with the screen at 100 cm) or accommodation (base-in prism with the screen at 20 cm). However, as with the TVOR, the magnitude of the effects showed considerable variability from one animal to another and, in some particular animals, from one direction to another.(ABSTRACT TRUNCATED AT 400 WORDS)     

Filling-in of visual phantoms in the human brain

The constructive nature of perception can be demonstrated under viewing conditions that lead to vivid subjective impressions in the absence of direct input. When a low-contrast moving grating is divided by a large gap, observers report seeing a 'visual phantom' of the real grating extending through the blank gap region. Here, we report fMRI evidence showing that visual phantoms lead to enhanced activity in early visual areas that specifically represent the blank gap region. We found that neural filling-in effects occurred automatically in areas V1 and V2, regardless of where the subject attended. Moreover, when phantom-inducing gratings were paired with competing stimuli in a binocular rivalry display, subjects reported spontaneous fluctuations in conscious perception of the phantom accompanied by tightly coupled changes in early visual activity. Our results indicate that phantom visual experiences are closely linked to automatic filling-in of activity at the earliest stages of cortical processing.     

Retinotopic maps and foveal suppression in the visual cortex of amblyopic adults

Amblyopia is a developmental visual disorder associated with loss of monocular acuity and sensitivity as well as profound alterations in binocular integration. Abnormal connections in visual cortex are known to underlie this loss, but the extent to which these abnormalities are regionally or retinotopically specific has not been fully determined. This functional magnetic resonance imaging (fMRI) study compared the retinotopic maps in visual cortex produced by each individual eye in 19 adults (7 esotropic strabismics, 6 anisometropes and 6 controls). In our standard viewing condition, the non-tested eye viewed a dichoptic homogeneous mid-level grey stimulus, thereby permitting some degree of binocular interaction. Regions-of-interest analysis was performed for extrafoveal V1, extrafoveal V2 and the foveal representation at the occipital pole. In general, the blood oxygenation level-dependent (BOLD) signal was reduced for the amblyopic eye. At the occipital pole, population receptive fields were shifted to represent more parafoveal locations for the amblyopic eye, compared with the fellow eye, in some subjects. Interestingly, occluding the fellow eye caused an expanded foveal representation for the amblyopic eye in one early–onset strabismic subject with binocular suppression, indicating real-time cortical remapping. In addition, a few subjects actually showed increased activity in parietal and temporal cortex when viewing with the amblyopic eye. We conclude that, even in a heterogeneous population, abnormal early visual experience commonly leads to regionally specific cortical adaptations.     

Afterimages: a collective term for percepts of different origin

Exposure of the eye to a strong photoflash results in a so-called "afterimage", which may last for 20 min or longer. In contrast, the true afterimage, which fluctuates in brightness and is best seen in complete darkness, lasts only a few minutes. This true afterimage can be attributed to the strong oscillatory neuronal responses immediately initiated by the flash. Thereafter dark and light regions, insensitivity percepts, are observable against bright and dark backgrounds, respectively. These percepts can be adequately explained by a reversal of the response behaviour of rod and cone driven ganglion cells situated along the contour of the flash-exposed area. The slow recovery of the rods explains why insensitivity percepts can be seen for many minutes.     

Eye-hand tracking using afterimages

Summary Oculomotor tracking of one's unseen hand is greatly enhanced when a positive afterimage of the arm is projected onto its changing apparent position. The afterimage arm also influences the perception of real arm motion. The nature of this influence indicates that sense of effort or will derives from spatial computations potentially involving sensory and motor information about the ongoing orientation of the entire body.Support was provided by NASA contract NAS 9-15147     

Accommodation and Size-Constancy of Virtual Objects

Accommodation has been suspected as a contributor to size illusions in virtual environments (VE) due to the lack of appropriate accommodative stimuli in a VE for the objects displayed. Previous experiments examining size-constancy in VE have shown that monocular cues to depth that accompany the object are a major contributor to correct size perception. When these accompanying cues are removed perceived size varied with the object’s distance from the subject, i.e., visual angle. If accommodation were the dominant mechanism contributing to a visual angle response [due to its action to keep physical objects clear] in this condition, an open-loop accommodation viewing condition might restore size-constancy to this condition. Pinhole apertures were used to open-loop accommodation and examine if size-constancy might be restored when few accompanying monocular cues to depth were present. Visual angle performance when viewing a low cue environment was found with and without the use of the pinhole apertures. Thus, these results signify that accommodation does not play a dominate role in the loss of size-constancy in sparse visual environments often used in VE. These results suggest that size-constancy is driven by the inclusion of the remaining monocular cues to depth in VE as it is in the physical world.