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Ehud Zohary

Publications and source records attributed to Ehud Zohary.

14 recordsLinked to original sources

Normal and abnormal fMRI activation patterns in the visual cortex after recovery from optic neuritis.

Recovery to normal or near normal visual acuity after an optic neuritis episode is common, despite frequent persistence of conduction abnormalities, evident in prolonged visual evoked potential (VEP) latencies. Improvement of visual function is commonly attributed to peripheral nerve recovery. However, central reorganization processes may also be involved. To assess this, we compared the patterns of fMRI activation, elicited by stimulation of the affected and the normal eye, along the visual cortical hierarchy. Activation was assessed in 8 subjects, which recovered clinically from an episode of optic neuritis but still had prolonged VEP latencies. In all patients, reduced fMRI activation was seen in V1 during stimulation of the affected eye, compared to the normal eye. The fMRI signal difference decreased in magnitude with progression along the visual hierarchy, and in some regions within the lateral occipital complex even showed the opposite preference (for the affected eye). These results may indicate a built-in robustness of the object-related areas to disruption of the visual input. Alternatively, it could reflect an adaptive functional reorganization of the cortical response to an abnormal input.

Adult↗

A mirror representation of others' actions in the human anterior parietal cortex.

The anterior part of the human intraparietal sulcus is known to be involved in visually guided grasping. This region is also active during the observation of object manipulation by others. Here, we explore the nature of action representation using functional magnetic resonance imaging (fMRI). Fourteen subjects observed video clips showing object manipulation by the right or left hand. The clips were presented in either the right or left peripheral visual field. The fMRI activation in the occipital cortex and in the caudal sections of the parietal cortex was specific to the visual-field location of the clips. In contrast, the principal factor determining the response in anterior intraparietal cortex was the identity of the observed hand. Furthermore, these "hand-specific" parietal areas also showed contralateral hand specificity during self action (i.e., object manipulation) without visual feedback. A similar selectivity for the identity of the observed hand was seen when using a region of interest analysis, focusing on individually defined visuomotor voxels within the parietal cortex. This dual visuomotor grasping representation lends further evidence for the existence of a mirror system in humans and suggests that the anterior intraparietal cortex is involved in the specific motor simulation of hand actions.

Adult↗

Visual cortex activation in bilingual blind individuals during use of native and second language.

Recent neuroimaging and transcranial magnetic stimulation studies indicate that the occipital cortex of congenitally blind humans is functionally relevant for nonvisual tasks. There are suggestions that the underlying cortical reorganization is restricted by a critical period. These results were based on comparison between early and late blind groups, thereby facing the problem of great variability among individuals within each group. Using functional magnetic resonance imaging, we studied bilingual congenitally blind individuals during use of 2 languages: one acquired early (Hebrew), the other later in life (English, at approximately 10 years). The subjects listened to chimeric words consisting of superimposed Hebrew and English nouns. They were instructed to either covertly generate a verb to the heard noun or repeat the noun, in either Hebrew or English. Lateralized activation during verb generation (vs. repeat) was found in classical language areas, in congruence with previous studies in sighted subjects. Critically, in our study, the blind participants typically also had robust left lateralized occipital differential activation during verb generation (vs. repeat), in both languages. This suggests that the critical period for plasticity persists beyond 10 years or that the visual cortex of the blind might be engaged in abstract levels of language processing, common to the 2 languages.

Acoustic Stimulation↗

Beyond retinotopic mapping: the spatial representation of objects in the human lateral occipital complex.

The spatial representation in the human ventral object-related areas (i.e., the lateral occipital complex [LOC]) is currently unknown. It seems plausible, however, that it would diverge from the strict retinotopic mapping (characteristic of V1) to a more invariant coordinate frame, thereby allowing for reliable object recognition in the face of eye, head, or body movement. To study this, we compared the fMRI activation in LOC when object displacement was limited to either the retina or the screen by manipulating eye position and object locations. We found clear adaptation in LOC when the object's screen position was fixed, regardless of the object's retinal position. Furthermore, we found significantly greater activation in LOC in the hemisphere contralateral to the object's screen position, although the visual task was constructed in a way that the objects were present equally often on each of the 2 retinal hemifields. Together, these results indicate that a sizeable fraction of the neurons in LOC may have head-based receptive fields. Such an extraretinal representation may be useful for maintenance of object coherence across saccadic eye movements, which are an integral part of natural vision.

Adult↗

Memory of ordinal number categories in macaque monkeys.

What mechanism underlies serial order memory? Studying preverbal serial memory shows that macaque monkeys reproducing a sequence of items can acquire knowledge of item ordinal position. In our previous experiment, macaques were repeatedly presented with image lists (first shown sequentially and then simultaneously on a touch screen together with a distractor chosen randomly from other lists). The task was to touch list images in the correct order. The monkeys' natural tendency was to categorize images by their ordinal position or number because their most common error was touching the distractor when it had the same ordinal number (in its own list) as the correct image. Item-to-item associations were used to complete the categorization strategy. Proposing a dynamic image-salience hypothesis for serial recall (based on category-to-image influence and a salience computation for identifying touch targets), we now study the category label characteristics in the context of this hypothesis. We found that these category labels are absolute, ordinal-number-based categories (first, second, etc.), not relative memorized as relative distance from the beginning and the end of the list, and not based on fixed ranking of reward contingency/image familiarity. Even isolated from item-item associations, the categories demonstrate category tuning (as well as the corresponding overlap of adjacent ordinal number codes). Moreover, monkeys choose images by proximity of their category to the current touch number, irrespective of the accuracy of the preceding choice. Category tuning itself is symmetric relative to correct ordinal position, but is skewed by other factors (reward, etc.). Tuning width increases with list length, with a concurrent increased use of item-to-item associations for determining touch order.

Animals↗

Dissociation between ventral and dorsal fMRI activation during object and action recognition.

Neuropsychological case studies suggest the existence of two functionally separate visual streams: the ventral pathway, central for object recognition; and the dorsal pathway, engaged in visually guided actions. However, a clear dissociation between the functions of the two streams has not been decisively shown in intact humans. In this study, we demonstrate dissociation between dorsal and ventral fMRI activation patterns during observation of object manipulation video clips. Parietal areas, such as anterior intraparietal sulcus (aIPS) display grasp viewing-dependent adaptation (i.e., fMR adaptation during repeated viewing of the same object-grasping movement) as well as a contralateral preference for the viewed manipulating hand. Ventral regions, such as the fusiform gyrus, show similar characteristics (i.e., adaptation, contralateral preference), but these depend on object identity. Our results support the hypothesized functional specialization in the visual system and suggest that parietal areas (such as aIPS) are engaged in action recognition, as well as in action planning.

Adult↗

V1 activation in congenitally blind humans is associated with episodic retrieval.

Recently we showed that the occipital cortex of congenitally blind humans is activated during verbal-memory tasks. Activation was found in regions corresponding to the retinotopic visual areas of sighted humans, including the calcarine sulcus (V1). No such occipital activation was found in sighted humans. One year later, the same blind subjects participated in a second fMRI scan, to study the contribution of semantic elements and episodic memory to the occipital activation. The subjects performed an episodic-memory task, requiring recognition of words that were originally presented in the first scan. We demonstrate here that the magnitude of V1 activation during the recognition task is correlated with memory performance, assessed during the scan. Across the blind, the better-remembered set of words elicited greater V1 activation than words from the poorly-remembered set, although the semantic components and the behavioral task were similar in the two sets. This indicates that on top of semantic processing (suggested previously), V1 activation in the blind is probably associated with long-term episodic memory. Indeed, within the blind, those who showed better recognition-memory performance had greater V1 activation compared with the poorer performers. We conclude that the posterior occipital cortex (including V1) of the congenitally blind is likely to be involved in episodic retrieval.

Adult↗

Transcranial magnetic stimulation of the occipital pole interferes with verbal processing in blind subjects.

Recent neuroimaging studies in blind persons show that the occipital cortex, including the primary visual cortex (V1), is active during language-related and verbal-memory tasks. No studies, however, have identified a causal link between early visual cortex activity and successful performance on such tasks. We show here that repetitive transcranial magnetic stimulation (rTMS) of the occipital pole reduces accuracy on a verb-generation task in blind subjects, but not in sighted controls. An analysis of error types revealed that the most common error produced by rTMS was semantic; phonological errors and interference with motor execution or articulation were rare. Thus, in blind persons, a transient 'virtual lesion' of the left occipital cortex interferes with high-level verbal processing

Analysis of Variance↗

Spatial vs. object specific attention in high-order visual areas.

Recently we reported that the topographic organization of visual field eccentricity in human visual cortex extends into high-order, ventral occipitotemporal (VOT) cortex. Within this cortex, regions that respond preferentially to faces and buildings have specific eccentricity biases, suggesting that this category-eccentricity association may reflect differential needs of recognition processes. However, it is still not clear to what extent this center/periphery differentiation within high-order occipitotemporal cortex depends on immediate, moment-to-moment, task demands. Previous attention studies were confined either to exploring the visual field topography (spatial attention) or to object identity (object-based attention). Here, we combined the investigation of these two different attentional mechanisms in the same study. We found that the main source of attentional modulation in occipitotemporal cortex was object-based attention. Shifting attention to different object categories (buildings, faces, and arrows) substantially modulated the object-related activations. The differential activation to each object category in occipitotemporal object areas was maintained, albeit at a reduced level, even when attention was focused on different spatial locations. A slight eccentricity-related attentional differentiation was observed in the more dorsal lateral occipital region, but not in the VOT cortex. These results argue against the possibility that the source of the eccentricity differentiation in VOT cortex is due solely to moment-to-moment shifts in spatial attention mechanism and supports the notion that the eccentricity-biased maps found in this region are due to built-in shape selectivity established over long-term processes.

Adult↗

Early 'visual' cortex activation correlates with superior verbal memory performance in the blind.

The visual cortex may be more modifiable than previously considered. Using functional magnetic resonance imaging (fMRI) in ten congenitally blind human participants, we found robust occipital activation during a verbal-memory task (in the absence of any sensory input), as well as during verb generation and Braille reading. We also found evidence for reorganization and specialization of the occipital cortex, along the anterior-posterior axis. Whereas anterior regions showed preference for Braille, posterior regions (including V1) showed preference for verbal-memory and verb generation (which both require memory of verbal material). No such occipital activation was found in sighted subjects. This difference between the groups was mirrored by superior performance of the blind in various verbal-memory tasks. Moreover, the magnitude of V1 activation during the verbal-memory condition was highly correlated with the blind individual's abilities in a variety of verbal-memory tests, suggesting that the additional occipital activation may have a functional role.

Acoustic Stimulation↗

Analysis of the neuronal selectivity underlying low fMRI signals.

BACKGROUND: A prevailing assumption in neuroimaging studies is that relatively low fMRI signals are due to weak neuronal activation, and, therefore, they are commonly ignored. However, lower fMRI signals may also result from intense activation by highly selective, albeit small, subsets of neurons in the imaged voxel. We report on an approach that could form a basis for resolving this ambiguity imposed by the low (mm range) spatial resolution of fMRI. Our approach employs fMR-adaptation as an indicator for highly active neuronal populations even when the measured fMRI signal is low. RESULTS: In this study, we first showed that fMRI-adaptation is diminished when overall neuronal activity is lowered substantially by reducing image contrast. We then applied the same adaptation paradigm, but this time we lowered the fMRI signal by changing object shape. While the overall fMRI signal in category-related regions such as the face-related pFs was drastically reduced for non-face stimuli, the adaptation level obtained for these stimuli remained high. We hypothesize that the relatively greater adaptation level following exposure to "nonoptimal" object shapes is indicative of small subsets of neurons responding vigorously to these "nonoptimal" objects even when the overall fMRI activity is low. CONCLUSIONS: Our results show that fMR-adaptation can be used to differentiate between neuronal activation patterns that appear similar in the overall fMRI signal. The results suggest that it may be possible to employ fMR-adaptation to reveal functionally heterogeneous islands of activity, which are too small to image using conventional imaging methods.

Adult↗

Convergence of visual and tactile shape processing in the human lateral occipital complex.

We have recently demonstrated using fMRI that a region within the human lateral occipital complex (LOC) is activated by objects when either seen or touched. We term this cortical region LOtv for the lateral occipital tactile-visual region. We report here that LOtv voxels tend to be located in sub-regions of LOC that show preference for graspable visual objects over faces or houses. We further examine the nature of object representation in LOtv by studying its response to stimuli in three modalities: auditory, somatosensory and visual. If objects activate LOtv, irrespective of the modality used, the activation is likely to reflect a highly abstract representation. In contrast, activation specific to vision and touch may reflect common and exclusive attributes shared by these senses. We show here that while object activation is robust in both the visual and the somatosensory modalities, auditory signals do not evoke substantial responses in this region. The lack of auditory activation in LOtv cannot be explained by differences in task performance or by an ineffective auditory stimulation. Unlike vision and touch, auditory information contributes little to the recovery of the precise shape of objects. We therefore suggest that LOtv is involved in recovering the geometrical shape of objects.

Acoustic Stimulation↗

Serial memory strategies in macaque monkeys: behavioral and theoretical aspects.

Serial memory is the ability to encode and retrieve a list of items in their correct temporal order. To study nonverbal strategies involved in serial memory, we trained four macaque monkeys on a novel delayed sequence-recall task and analysed the mechanisms underlying their performance in terms of a neural network model. Thirty fractal images, divided into 10 triplets, were presented repeatedly in fixed temporal order. On each trial the monkeys viewed three sequentially presented sample images, followed by a test stimulus consisting of the same triplet of images and a distractor image (chosen randomly from the remaining 27). The task was to touch the three images in their original order, avoiding the distractor. The monkeys' most common error was touching the distractor when it had the same ordinal position (in its own triplet) as the correct image. This finding suggests that monkeys naturally categorize images by their ordinal number. Additional, secondary strategies were eventually used to avoid distractor images. These include memory of the sample images (working memory) and associations between triplet members. Further direct evidence for ordinal number categorization was provided by a transfer of learning to untrained images of the same ordinal category, following reassignment of image categories within each triplet. We propose a generic three-tier neuronal framework that can explain the components and complex set of characteristics of the observed behavior. This framework, with its intermediate level representing ordinal categories, can also explain the transfer of learning following category reassignment.

Animals↗

Contrast sensitivity in human visual areas and its relationship to object recognition.

An important characteristic of visual perception is the fact that object recognition is largely immune to changes in viewing conditions. This invariance is obtained within a sequence of ventral stream visual areas beginning in area V1 and ending in high order occipito-temporal object areas (the lateral occipital complex, LOC). Here we studied whether this transformation could be observed in the contrast response of these areas. Subjects were presented with line drawings of common objects and faces in five different contrast levels (0, 4, 6, 10, and 100%). Our results show that indeed there was a gradual trend of increasing contrast invariance moving from area V1, which manifested high sensitivity to contrast changes, to the LOC, which showed a significantly higher degree of invariance at suprathreshold contrasts (from 10 to 100%). The trend toward increased invariance could be observed for both face and object images; however, it was more complete for the face images, while object images still manifested substantial sensitivity to contrast changes. Control experiments ruled out the involvement of attention effects or hemodynamic "ceiling" in producing the contrast invariance. The transition from V1 to LOC was gradual with areas along the ventral stream becoming increasingly contrast-invariant. These results further stress the hierarchical and gradual nature of the transition from early retinotopic areas to high order ones, in the build-up of abstract object representations.

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